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OCCASIONAL PAPERS 


OF THE 


MUSEUM OF ZOOLOGY 


UNIVERSITY OF MICHIGAN 


NUMBERS 129-152 
VOLUME VI 
1923-24 


ANN ARBOR 


PUBLISHED BY THE UNIVERSITY 


1 salou ea L 
ADVERTISEMENT 


The publications of the Museum of Zoology, University of 
Michigan, consist of two series—the Occasional Papers and the 
Miscellaneous Publications. Both series were founded by Dr. 
Bryant Wall Myr. Bradshaw H. Swales and Dr. W. W. 


Papers, publication of which was begun 

iedium for the publication of brief original 

principally upon the collections in the Museum. 

re issued separately to libraries and specialists, 

icient number of pages have been printed to 

tle page, table of contents, and index are 

up} libraries and individuals on the mailing list for 


ous Publications inelude papers on field and 


ue nographic studies and other papers not 

ithin the scope of the Occasional Papers. The papers are 

published separately, and, as it is not intended that they shall 

be grouped into volumes, each number has a title page and 
table of contents. : 


ALexanpEr G. RuTHVEN, 
Director of the Museum of Zoology, 
University of Michigan. 


i 


TABLE OF CONTENTS 


No. 129. Barbour, Thomas.—Notes on Reptiles and Amphibians from 
Panama. 

No. 130. Williamson, E. B.—Notes on the Habitats of Some Tropical 
Species of Hetaerina (Odonata). 

No. 131. Barbour, Thomas.—The Crocodile in Florida. 

No. 132. Barbour, Thomas.—West Indian Investigations of 1922. 

No. 133. Chamberlin, Ralph V.—Results of the Bryant Walker Ex- 
peditions of the University of Michigan to Colombia, 1913, and 
British Guiana, 1914. The Diplopoda. (29 plates.) 

No. 134. Williamson, E. B.—A New Species of Archaeogomphus 
(Odonata). (1 plate.) 

No. 135. Baker, H. Burrington.——The Mollusca Collected by the Uni- 
versity of Michigan-Walker Expedition in Southern Vera Cruz, Mex- 
ico, IV. (1 plate.) 

No. 136. Ruthven, Alexander G., and Helen T. Gaige.—Description of 
a New Species of Pipa from Venezuela. 

No. 137. Baker, H. Burrington—The Mollusea Collected by the Uni- 
versity of Michigan-Williamson Expedition in Venezuela. Part I, 
Curacao. (5 plates.) 

No, 138. Becker, Herbert Ray.—The Habitat of Aphredoderus sayanus 
in Kalamazoo County, Michigan. 

No. 139. Ortenburger, Arthur I—A Note on the Genera Coluber and 
Masticophis, and a Description of a New Species of Masticophis. (3 
plates. ) 

No. 140. Blanchard, Frank N.—A New North American Snake of the 
Genus Natrix. (1 map.) 

No. 141. Dodds, G. S—A New Species of Phyllopod. (1 plate.) 

No. 142. Blanchard, Frank N.— Comments on Ring-neck Snakes 
(Genus Diadophis), with Diagnoses of New Forms. 

No. 143. Ruthven, Alexander G.—The Reptiles of the Dutch Leeward 
Islands. 

No. 144. Hubbs, Carl L.—A Note on the Species of Evermannichthys, 
a Genus of Sponge-Inhabiting Gobies. 

No. 145. Winslow, Mina L.—Two New Freshwater Snails from Mich- 
igan. (1 plate.) 

No. 146. Koelz, Walter—Two New Species of Cisco from the Great 
Lakes. 

No. 147. Ruthven, A. G., and Helen T. Gaige—A New Leposoma 
from Panama. ~ 

No. 148. Hubbs, Carl L.—Studies of the Fishes of the Order Cyprino- 
dontes. V. Notes un Species of Goodea and Skiffia. 

No. 149. Ruthven, Alexander G.—Description of an Ameiva from 
Testigos Island. 

No. 150. Blanchard, Frank N.—A New Snake of the Genus Arizona. 

No. 151. Dunn, E. R.—Some Panamanian Frogs. (2 plates. ) 

No. 152. Baker, H. Burrington—lLand and Freshwater Molluses of 
the Dutch Leeward Islands. (21 plates.) 


ill 


NUMBER 129 JANUARY 25, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


~ 


NOTES ON REPTILES AND AMPHIBIANS FROM 
PANAMA 


By THomaAsS BARBOUR 


This spring Mr. W. S. Brooks and I decided to make a 
collection of vertebrates in the hitherto unvisited Eastern Sapo 
highlands. From this serrania flow many of the tributary 
streams of the Sambi River in eastern Panama. Unusually 
early and heavy rains made this a task of more than ordinary 
difficulty. Our Chocoano Indian companions were anxious 
to plant their depleted gardens, rather than to toil as carriers 
in the stifling forest, and it was with considerable difficulty 
that they were persuaded to help us. Their consent to our 
plans once obtained, they proved loyal and useful allies, and 
our fruitful journey ‘is the direct result of their aid. Late 
one afternoon in early March we were dropped off at the 
head of navigation on the Sambi River, just where the Sabalo 
enters this stream fro: the south. We had left Panama City 


on board the Panamanian schooner “Chiriqui,” then under 


2 University of Michigan 


government charter, since a chance official excursion to settle 
a disputed oil claim on the coast afforded us this welcome 
opportunity to reach the little fever-ridden Panamanian coast 
settlement of Garachiné near the Sambi River mouth. 
Thanks to Dr. R. P. Strong, who accompanied us as far as 
the mouth of the river, we got up the stream to Boca de 
Sabalo in a small motor launch towed by the schooner from 
Panama. After a long colloquy with the Indians, we decided 
to strike north to a range of hills visible from our landing 
place. Several hours’ walk over a rough trail brought us to 
the Rio Esnape, evidently a tributary of the Rio Taimiti, and 
one which rose in the hills now near at hand. Here we col- 
lected several days, but the rains became constant and it was 
almost impossible to dry our collections of bird and mammal 
skins, and besides, reptiles were very scarce. We then 
recrossed the Sambu and marched overland southward to the 
head of the Rio Jesus, which enters the Sambi far down- 
stream from the mouth of the Sabalo and which runs parallel 
to the Sabalo, but arises much farther inland. Several camps 
were established along this little river, and then we marched 
to the upper Rio Jesusito, a swift torrent, said by some to 
be a source stream of the Rio Celorio. The latter small river 
enters the sea ;just south of the mouth of the Sambu, nearer 
Garachiné point.t_ We made two camps in the foothills of the 
Sapo range of mountains along the Alto Jesusito. We then 
moved over to the headwaters of the Rio San Antonio and 
established a camp on the slopes of the Cerro de Sapo, a peak 
nearly 2,000 meters in altitude. From this point we carried 


down to Garachiné again and returned in a twenty-two-foot 

1Other Indians declared that the Jesusito “has no mouth,” so we 
inferred that it might find its way into some great marsh and not 
really connect with the Celorio at all. 


Occasional Papers of the Museum of Zoology 3 


motor launch—not without various vicissitudes—to Panama. 
The novelties now described have appeared during an exam- 
ination of the reptiles and amphibians caught during this jour- 
ney. Other notices will appear later dealing with the mam- 


mals, birds, and fishes. 


The region is one of high, damp, humid forest, gloomy 
and stifling except where some water course cuts through the 
wooded lowlands, letting in the sunlight. Decay of fallen 
wood and leaves is very rapid and the dark forest floor is 
sodden and slippery. In general, reptiles were surprisingly 
rare, and often a day would pass when none of us would see 
a lizard, unless when coming to the shore of some small 
stream the bipedal basiliscs would scurry away. The young 
far outnumber the adults and all are well able to run with 
equal ease over land or the face of the water. While running 
the body is held almost upright, the tail is raised as a balance, 
and the fore limbs are tightly pressed to the sides. ‘They 
move and stop with a speed and precision which seems mechan- 
ical rather than animate. The paucity of adults and the shy- 
ness of both young and old bespeak abundant enemies, but of 
what nature we were never able to learn. 

One afternoon an Indian who had been gathering firewood 
came in carrying a small lizard, and we then saw for the first 
time the young of Diploglossus monotropis, already known 
from Costa Rica and Colombia. This little creature, about 
seven inches long, was so gorgeously colored while alive and 
so different from the preserved examples that my field notes 
are worth copying. “This specimen, seven inches long, has a 
grey-green head, brilliant carmine sides covered with anasto- 
mosing black lines; belly yellowish; back and tail black with 


beautiful narrow blue-grey, almost mauve cross-bars.” I have 


4 University of Michigan 


neer seen such a splendid lizard except my Duploglossus 
resplendens from Bolivia. 

A few days after our Upper Jesusito camp was made we 
began to fell trees to let in sunlight and breeze. As it turned 
out, there was no breeze and the sun was almost constantly 
obscured by rain clouds. One tree came down with a crash 
and brought with it a living and uninjured Corythophanes cris- 
tatus. ‘The interesting point in connection with this capture 
was the fact that we chanced to keep the lizard alive long 
enough to find that its actions were singularly chamaeleon-like. 

It was sluggish and deliberate in its movements, and when 
angered it reared upright, flattened its body vertically, and 
bent down its head. Its mouth meanwhile was opened widely 
in a way that recalled at once captive and angry African 
Chamaeleons. That the very peculiar superficial similarity of 
appearance should be accompanied by such similar sluggish 
movements and curious attitudes is most noteworthy and 
almost incredible when the protean zoologic gap between the 
two genera is considered. 

In a few places where the forest roof leaked spots of sun- 
light the ground did dry out and the great, curly, new-fallen 
leaves made noisy walking. In these little dried out spaces 
we found some tiny lizards. They crept swiftly and stealthily 
over the big dead leaves, and when the sun was hidden, as it 
often was because of the frequent showers, these little lizards 
hid at once, to reappear when their mouldy abode became dry 
again. They were not easy to catch, and when one was finally 
in the fingers a decent specimen was by no means assured, 
for their skin tore like wet tissue paper and their struggles 
usually left them sadly unfrocked. 

These, as other slim-toed gekkos or Eublepharids, as they 
once were called, are far more agile than their allies with 


Occasional Papers of the Museum of Zoology 5 


dilated digits—more alert and less deliberate in their move- 


ments.. When examined the species proved to be the rare 


and little known Lathrogecko sanctae-martae Ruthven. 

Their color is very characteristic and varies but little. 
They are rich mahogany brown above and grey below. A 
narrow, light lateral line decorates each side and is most con- 
spicuous on the posterior half of the trunk and fades on the 
basal third of the tail-or does not extend so far. A conspicu- 
ous light marking shaped like a horseshoe encircles the occi- 
put and two white lines extend from in front of the eyes to 
meet on the tip of the snout. The belly is immaculate, the 
tail reticulated and speckled below. 

_ A considerable series was secured. The head scales afford 
poor diagnostic features. 

The woodland Anoles were rare. One day brought to bag 
what is probably the giant Anolis latifrons Bertholdt. This 
lizard has a great apple-green throat fan with small, dark 
spots and is a striking creature in life. Previously known 
only by the type from Popoyan, it is easy to see how Ber- 
thold’s figures may have misled Boulenger. The drawing 
(Verh. Ges. Gottingen 3, 1847, p. 6, pl. 1, fig. 2), where it is 
not frankly diagrammatic, is fairly accurate, so far as the 
topography of the cephalic shields is concerned, but the sculp- 
turing is apparently not drawn in. In reality the head scales 
are rougher than in A, squamulatus, instead of the reverse as 
stated by Boulenger (Cat. Lizards B. M., 2, 1885, p. 62). 
The real character easily separating the two species is the 
presence in Jatifrons of a series of enlarged tubercle-like scales 
along the supraocular margin. In squamulatus, which we did 
not take, but which is well represented in the U. S. National 
Museum, the small granular scales extend from the supra- 
ocular disc to the very edge of the area over the eye, quite 


6 University of Michigan 


without the development of any marginal tubercles. This ill- 
drawn figure—and the description is also perfectly vague— 
apparently misled Boulenger into re-naming Anolis latifrons 
as Anolis princeps.2 I have seen an authentic specimen of this 
form in the University of Michigan Museum, thanks to Dr. 
Ruthven. 

The most.common species met with was what we con- 
sider to be Anolis limifrons, a slender little beast with an 
ivory white dewlap in life, and Anolis binotatus, reaching a 
far larger size and with the dewlap brilliant carmine enclosing 
a small, black, central spot. In life binotatus is rich ‘olive- 
greenish, with dirty yellow blotches on the limbs and dark 
markings on the body. It has a way of resting rather flattened 
against bark with its legs sprawling and spread out, and the 
whole result of attitude and coloration is very strikingly 
lichen-like. The narrow dark band between the eyes and the 
dark perioccipital vitta are conspicuous and constant mark- 
ings in life. We secured a few specimens of Anolis stig- 
mosus, but the species seems much less abundant on the main- 
land than on the islands in the Gulf of Panama. We only 
found one other true Anolis which was a rather common spe- 
cies, having a dewlap pinkish toward the margin and dusky 
toward the base, if I remember correctly. Unfortunately, I 
omitted to make notes of this in life. After many perplexi- 
ties, I have concluded that this form is so near Anolis gaiges 
Ruthven of the Santa Marta Mountains in Colombia that for 
the present, at any rate, our Sapo Mountains individuals had 
best bear this title. 

Our most surprising treasure was a new genus of rather 
baffling affinities. ‘The form of the body and the structure of 
the digits recalls true Anolis; the cephalic squamation resem- 


2Ann. Mag. N. H., (7), 9, Jan., 1902, p. 54. 


Occasional Papers of the Museum of Zoology 7 


bles that of Xiphocercus, or somewhat less so of Phenaco- 
saurus, while the scales of the body and the form and struc- 
ture of the gular appendage—it is never a “pouch’—are strik- 
ingly unlike any of the Anolinae now known. 

It is important also to remember that the dewlap of Tropi- 
dodactylus, which Boulenger states is “not inflatable,” is really 
capable of being both contracted and folded so that it is 
decidedly similar to the dewlap of Anolis, which can also be 
expanded, but never, of course, “inflated.” Tropidodactylus 
represents an Anoline stock which has become modified for 
fossorial life as Norops is for a purely cursorial existence. 


The new genus may be called 


Diaphoranolis gen. nov. 


Tympanum distinct; body very strongly compressed, 
entirely covered with juxtaposed pavement-like scales; no 
dorso-nuchal crest; male with a non-extensible, plicate, pen- 
dulous gular appendage covered with scales similar to those 
of the body; digits evenly and rather extensively dilated, with 
subdigital lamellae, the distal joints slender and raised as in 
Anolis ; no inguinal pores ; tail long, much compressed ; it curls 
laterally and probably is slightly prehensile as in some Anoles, 
notably homolechis; lateral teeth tricuspid; abdominal ribs 


present. 


Diaphoranolis brooksi® sp. nov. 


Type: M. C. Z. 16,297, from Mt. Sapo, eastern Panama, 
2,500 feet elevation; Barbour and Brooks, collectors; April, 
1922. Head medium, nearly twice as long as broad; forehead 
very slightly concave, covered with rather large, irregular 


3’ Named for my friend and frequent companion, Winthrop Sprague 
Brooks, Esq., of the Boston Society of Natural History. 


8 University of Michigan 


pavement-like flat shields; supraorbital semicircles each com- 
posed of two rows of shields, the inner series of each semi- 
circle separated from its fellow of the opposite side by two 
rows of small, irregular shields; occipital scale of irregular 
oblong shape, about as large as ear opening, separated from 
the semicircles by three series of rather large plates; supra- 


ocular disks large, composed of eight or ten enlarged shields 
in contact with scales of gradually diminishing size in the other 


directions ; canthus rostralis rather obtuse, but sharply defined 
by a straight linear suture separating the enlarged canthal 
shields and the almost equally enlarged shields of the upper 
loreal row; four loreal rows below this enlarged upper series; 
the lowest row, however, bordering the supralabials, being 
also much enlarged; eleven supralabials, nine infralabials; 
gular appendage large, pendulous, plicate, and covered with 
non-imbricating; pavement-like scales similar to those on the 
rest of the body; body very strongly compressed, slender and 
delicately formed, covered entirely with flat non-imbricating, 
plate-like scales, so that all scales of head and body are undif- 
ferentiated in their essential features, except that those of 
the head are larger and polygonal, whereas on the rest of the 
body they are rather small, more or less similar in size, except 
that the ventrals are larger than either the laterals or the dor- 
sals; the dorsals tend to be round, the laterals are all more or 
less oblong and the ventrals tend to be\|hexagonal in shape; 
the limbs are rather short, the adpressed hind limb reaches 
just anterior to the shoulder; about forty lamallae beneath the 
fourth toe; tail strongly compressed, the basal row of plate- 
like scales on each side keeled, all the rest smooth. 

Color in life: Very light china bluish-grey, changing to 
white in alcohol, the head, neck and dewlap all similarly 


marked with a network of coarse black lines, two black sad- 


Occasional Papers of the Museum of Zoology 9 


dles on the back and nine black rings completely surrounding 
the tail; limbs and digits with many sharply defined pairs of 
fine, black lines occurring as rings which do not quite meet 
on the inner surface. 

Total length of head and body, 132 mm.; tail, 86 mm.; 
head, 15 mm. 

This most curious and strikingly colored lizard is now 
pallid white, with many black markings very sharply defined. 
The absence of flash colors from the pendulous dewlap and 
the extension out to the very margin of this appendage of the 
same network of markings that are so unique and surprising 
a feature of the head and neck is wholly unexpected. The 
dewlap appears more like that of Iguana in miniature. I do 
not believe that this type can be considered more primitive 
or more advanced than any of the other allied genera. Some 
of the Anoline genera may well have sprung from species of 
Anolis itself as we know it, but it seems probable that this 
genus sprung from some common ancestor. The expanding 


dewlap with a flash-color is obviously desirable, for it is wide- 
spread and shows large variety of development within the 


great genus Anolis itself. That type of dewlap presupposes 
a stylus and muscles to make expansion possible, and the nec- 
essary accommodation can only be obtained with imbricating 
scales on rubbery, elastic skin. The appendage in this genus, 
obviously also a development for purposes of ornamentation, 
accomplishes its more modest attempt at beautification in a 
wholly different manner. 

Along the stream beds the diurnal and nocturnal fauna— 
as might be expected—differed widely. Ameiva undulata 
quadrilineata and Ameiva festiva, the latter with a sky-blue 
mid-dorsal stripe in the young and looking most scinc-like, 
were about equally abundant. Bufos were common, typhonius 


fe) University of Michigan 


and haematiticus along the woodland streams; the former 
diurnal and the latter nocturnal; marinus hopped about all 
the clearings, and a rare and rather atypical sternosignatus 
was found sparingly in the woods. Eupemphix pustulosus 
was likewise a diurnal denizen of the shady forest and many 
were found. Eleutherodactylus palmatus we took but once, 
and that by day. These two species were caught among the 
scattered stones and pebbles of the shores and beaches of the 
river near our camp. Eleutherodactylus ranoides we found 
several times, but with the night lamp only. Leptodactydac- 
tylus melanonotus was diurnal and common; the others taken 
were found only at night. Night hunting with the light, we 
often caught the great L. pentadactylus, and as our containers 
did not permit of our carrying off large series we had ample 
opportunity to test the remarks of good Father Labat, who 
remarked in 1724, when he first tasted them in Guadeloupe, 
that they were “les plus belles Grenonilles du Monde.” By 
day they keep hidden in their caves and are never seen. They 
emerge at night and sit in the shallows awaiting their prey— 
usually shrimps. Leptodactylus bolivianus, which I once 
redescribed as L. insularum, was likewise often caught, and 
only after dark, unless we chanced to dig it from its deep and 
almost perpendicular burrow. Rana palmipes for some reason 


seemed to be distinctly rare and we only found it once. 


Four species of tree toads were taken during the trip, and 
four only. All these were found with the aid of a hunting 
lamp at night. Hyla maxima Laur. is apparently new to the 
fauna of Panama and was found at the Rio Esnape. The 
second species is represented by a young individual too small 
for satisfactory identification. The third I consider worthy 


of subspecific recognition. 


Occasional Papers of the Museum of Zoology II 


Hyla baudinii dolomedes, sp. nov. 

Type: M. C. Z. 2,39, from the Rio Esnape, Sambu Valley, 
eastern Panama. Barbour and Brooks, 1922. 

Similar to true H. baudinii of Central America, but with 
very long hind limbs—longer than Central American individ- 
uals which I have seen. The tibio-tarsal articulation reaches 
well beyond the tip of the snout. 

Boulenger (P. Z. S., 1913, p. 1,023) remarks: “One of 
the specimens (from the Colombian Choco), a female, is 
remarkable for the longer hind limbs, the tibio-tarsal articu- 
lation reaching beyond the tip of the snout.’ We are not 
informed whether the other individuals may not also have 
had limbs longer than normal, if less strikingly so. In any 
case, such frogs seem unknown in upper middle America, and 
even if there is overlapping as well as possible intergradation 
between the form it is worthy of a name for convenience. The 
form is probably worth full specific recognition. 

The fourth species found, I believe, represents Dr. Noble’s 
Hyla chica, The three examples taken agree fairly well with 
a paratype of chica, which seems to have a vastly greater 
range than one would expect for such a tiny form. 


The species of Atelopus are-at best but half known. Afe- 
lopus varius as now understood has a very great range, and 
wherever it has been collected in numbers it appears to vary 


greatly both individually and geographically. 


This spring we camped for a week or more by a small 
stream, one of the headwaters of the Rio San Antonio on the 
slopes of Mt. Sapo. Little frogs of the genus Atelopus were 
common and we observed them daily. Singularly lethargic, 
they were usually perched on some projecting stone in mid- 
stream, and when disturbed they flopped feebly into the 


12 University of Michigan 


water and were carried down the brook, striking out lazily 
until they reached some chance refuge. They were easily 
caught, and we preserved nearly fifty examples. This series 
shows almost no variation in color, also no marked structural 
differences from varius, but it lacks the inherent quality of 
varius, which is variability. A. spurrelli Boulenger (P. Z. §$., 
1914, p. 813, pl. 1, fig. 1) is probably related, but is known, 
however, from only a single specimen, so it is impossible to 
do more than suggest that possibly the A. varius stock may 
have given rise to some local types in the great Chocoan forest 


region in which a fixity of color pattern has been attained 
The coloration, too, is unique and striking. 


Atelopus spurrelli certus, subsp. nov. 
Type: M.C. Z. 8,538, from a stream on Mt. Sapo, east 
ern Panama. Barbour and Brooks, 1922. Paratypes in M. 
CoZ. and the A. M,N; H- 


Similar to A. s. spurrelli as described and figured by Bou- 
lenger, but with the dark dorsal areas broken up into series 
of spots and blotches, with, nevertheless, the underlying 
topography being preserved in almost every case. 

The belly has a tendency to be more finely spotted in the 
males and almost immaculate in the females; in which sex 
also the dorsal dark patches are much more finely comminuted 
than in the males. 

The creature is brick red, almost vermillion, in life, with 
the dark spots velvety black. In alcohol this ground color 
has faded to whitish on the belly and to a pallid reddish hue 
on the light dorsal interspace between the black blotches which 
have remained essentially unchanged. 

Noble, after examining some of our series, declared them 
to be A. varius pure and simple, and we marvel at our own 


Occasional Papers of the Museum of Zoology 13 


temerity in not at once accepting his verdict. There is, how- 
ever, the question of whether possibly spurrelli is not really 
as distinct as Boulenger considered it to be, and no one can 
doubt the close affinity of spurrelli and certus. For the pres- 
ent, therefore, it seems wisest to keep these various catagories 
separate, although there is no doubt but that very possibly 
adequate material might reduce both these names to be syno- 
nyms of varius. 

The common Dendrobates tinctorius so abundant in the 
dry woods on Ancon hill in the Canal Zone and the islands, 
especially Taboga, near the Pacific mouth of the canal, differ 
conspicuously from the individuals from eastern Darien. The 
Canal Zone poison-toads are rich velvety black or more rarely 
very deep purplish maroon, with large, irregular blotches of 
the most vivid metallic green. The specimens from the Sambu 
Valley have the same dark ground color, but differ in always 
having the vivid green occur as small round dots about one- 
eighth inch in diameter and widely scattered. Such a host 
of color phases of D. tinctorius have been noticed in the liter- 
ature that, until the far day arrives when material is assem- 
bled for an adequate revision, it is unwise to name the one 
in hand. Future collecting, to be useful, must best be done 
by the reviser himself that he may see in life these creatures 
which pass through such instant metachroses in alcohol. 

To sum up, we may say that in general in this damp rain 
forest reptiles are surprisingly few. Amphibians, too, are far 
from being the abundant creatures which they often are in 
other similar situations. We saw but two small snakes during 
all our tramps, and both of these belonged to the common 
Leptodeira annulata. One fell from a small tree we cut down; 
the other, one of the men killed while night hunting. About 
the Canal Zone snakes were far more abundant. 


14 University of Michigan 


Thanks to Mr. James Zetek and Dr. Clark, both connected 
with the Board of Health Laboratory at Ancon, we were not 
only able to have specimens brought in to the laboratory and 
preserved during our absence in Darien, but were given a 
number of specimens which from time to time had been 
brought to the laboratory for examination, \usually to deter- 


mine whether or not they were venomous. 


Among the rarities so found by Brooks and myself were 
Coecilia sabogae Barbour, previously only known from the 
Pearl Islands. This specimen agrees well with the type and 
has 14 or I5 vomerine teeth, eight or nine teeth on each 
side of the upper jaw; eight teeth on each side of the outer 
row of the lower jaw and three on the inner row. The four 
anterior maxillary and mandibular teeth are much enlarged. 
This species is now to be recorded from the vicinity of Ancon. 

Among the snakes, Micrurus nigrocinctus (Gir.) and its 
remarkable counterpart, Erythrolamprus aesculapu (Linné), 
caught within a few days of each other in almost the same 
spot and under similar conditions, offered a most graphic 
exposition of this ill-explained phenomenon of “mimicry.” In 
any case, whatever may be the cause of the coloration, in both 
species the similarity is almost certainly purely fortuitous. 
Himantodes elegans (Jan.), known from Costa Rica and Gua- 
temala, occurred with H. cenchoa (Linné) about Ancon. Lep- 
tocalamus torquatus Giinther is another rare and little known 
species represented by two specimens in our Ancon collection. 
The ten other species secured were all common and already 
well known from the locality. 

One novelty has, however, appeared, a single Micrurus, 
and one which has been carefully examined by my friend, 


Dr. Dunn, who is particularly interested in this genus. It is 


Occasional Papers of the Museum of Zoology 15 


closely related to both M. tschudii (Jan.) and M. dissoleucus 
Cope. It may well be called 


Micrurus dunni, sp. nov. 


Type: M. C. Z. 16,304, from the vicinity of Ancon, Canal 
Zone of Panama. 

Head only moderately depressed; diameter of eye equal 
to about two-thirds of its distance from mouth; rostral much 
broader than high, little visible from above ; frontal very small, 
about equal to a supraocular in breadth, not twice as long as 
broad, and much less than its. distance from the tip of the 
snout; one prae- and two postoculars; anterior temporal pres- 
ent and very narrow; seven upper labials, third and fourth 
entering eye and nearly equal in size, the seventh well devel- 
oped; first lower labials in contact behind the symphysial ; 
parietals very large, longer than their distance from the inter- 
nasals; scales in 15 rows; anal divided; ventrals, 224; sub- 
caudals, 109. 

Color: Head black, with a narrow white band crossing 
the anterior portion of the parietals and sending forward 
extensions to the posterior border of each eye, to cover two- 
thirds of the frontal and the fifth and half of the sixth labial; 
this white ring is followed by a wide black band, then a nar- 
rower red band, and then eleven triads, on the body, in each 
case the central black ring being the widest, the other two 
black bands being equal in width to the red interspaces sepa- 
rating the triads and the same outer black rings of each triad 
about twice as wide as the red rings within the triad; these 
red rings also have many scales tipped with black; tail with a 
single normal triad and a black tip. 

Total length, 130 mm.; tail, 20 mm.; diameter of body, 
4.5 mm. 


16 University of Michigan 


Micrurus hollandi (Griffin), a species recently described 
and one which seems to have been missed in compiling the 
Zoological Record, belongs also to this section of the genus. 
It is, however, distinct from the one now described and came 
from Bonda, Colombia. (Mem. Carnegie Mus., Pittsburg, 7, 


3, 1915, p. 218.) 


NUMBER 130 FEBRUARY I0, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arzor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


NOTES ON THE HABITATS OF SOME, TROPICAL 
SPECIES OF HETAERINA (ODONATA) 


By E. B. WILLIAMSON 


GENERAL CONSIDERATIONS 


Dragonflies are comparatively rare in nature, although 
many sometimes congregate in a very restricted habitat, as a 
narrow belt of vegetation about a lake or the ripple on some 
stream, and, from observations at such stations, one may get 
an exaggerated idea of the number of individuals of certain 
species. The activity of some species often gives the same 
impression. A small pond near Nirgua, Venezuela, one day 
was “alive with Trameas.” Late that afternoon the Trameas 
abandoned the pond to rest a few feet from the ground on 
the twig tips of nearby dead bushes. Lowering one’s head 
to the ground brought these resting dragonflies against the 
sky line and it was an easy matter to see and net all these 
resting individuals which had successfully eluded the net dur- 
ing the day. A few trips around the pond resulted in the 


2 University of Michigan 


capture of all of them, and the total was less than thirty indi- 
viduals. The next day at the pond Trameas were conspicuous 


by their absence. 


Once in Guatemala I had collected about Gualan for sev- 
eral days with poor success. It was the very end of the dry 
season, and the woods and fields were tinder dry. Then one 
night it rained, a veritable downpour, for hours. And about 
the little ponds which came into being between sunset and 
sunrise along the railroad embankment Trameas and other 
libellulines flew “by hundreds” in the early morning sun. And 
yet, during the preceding days of drought I had not seen a 
Tramea. They were really few in numbers and were scat- 
tered far and wide through the brush and over the fields. 
Dragonflies would be almost unknown except to a few spe- 
cialists, if it were not for their congregating at times of 
greater or lesser duration in habitats of very restricted area. 

Yet these habitats may be occupied for only a brief period 
in the lives of the dragonflies. Because of their freedom of 
flight, their relative independence of any one certain food, 
and their limited numbers, dragonflies present in many cases 
a difficult problem for one who would discuss their habitats 
intelligently. To say, for example, of some species of Soma- 


I 


tochlora that it “frequents woodland streams,” may tell about 
as much about it as the student would learn of the activities 
of Charles Darwin, say, if his biographer told where Mr. 
Darwin spent his youth and gave the street address and house 
number of his later years. For the chances are that our 
Somatochlora from the date of its emergence till its final 
activity (mating) may never visit the woodland stream. We 
find it at a certain period in its life at a habitat of very small 


area, where it meets others of its kind. 


Occasional Papers of the Museum of Zoology 3 


Dragonflies as adults, when they are usually most readily 
observed, may have a very short life, and a few days differ- 
ence in visiting some suitable habitat may result in the failure 
to find a single individual of a certain species which may have 
been there in numbers a few days before or after the visit. 

Some dragonflies also vary from year to year, due to early 
or late seasons, in the time of their appearance. If their sea- 
son is very short, the student, if not continuously in the field, 
looking for the species only on certain dates, may miss it 
entirely. 

Some dragonflies, if not all, seem to occur in relatively 
large numbers only at intervals of undetermined duration. 
The species maintains itself continuously in a certain habitat, 
but has “lean and fat years.” This phenomenon may be 
obscured or confused by the ebb and flow of odonate life 
described in a later paragraph. 

Many dragonflies are very susceptible to some other con- 
ditions which are probably meteorological. On a certain day 
a pond, for example, may be alive with the active individuals 
of many species. The succeeding day may be, to the observer, 
similar to the day before, but at the same pond fewer species 
and a smaller number of individuals may be on the wing. 

The student of habitats encounters another difficulty. 
Dragonflies “come and go’—there are great ebbs and flows 
over long periods. I do not refer to the occasional individual 
strays or waifs, of which the collector finds a few during 
many years of collecting, but rather to large and, apparently 
at the time, successful invasions of a new habitat. Two good 
examples come to mind. Near Bluffton, Indiana, is a wood- 
land swamp of possibly three or four acres. I have had this 
pond under observation for over twenty years. One year 


Inbellula quadrimaculata appeared there in great numbers. 


4 University of Michigar 


They were easily the dominant dragonfly and any student of 
habitats would have pronounced them one of the essential 
factors in the balance of this woodland pond. But Libellula 
quadrimaculata has appeared at this pond only once. Twice 
during my observation of this pond Enallagmas have appeared. 
One of these years was 1920, when Enallagmas probably 
exceeded in numbers all other dragonflies on the wing at the 
pond during the same season. Flying a few inches above the 
water, resting on grass stems, willow tips, and spatter-dock 
leaves, on the broad surfaces of which the females were ovi- 
positing, they dominated the entire pond. But they were not 
there in 1919 or in 1921. The changes which take place from 
year to year in any odonate society such, for example, as an 
abandoned gravel pit, may be obscured or subject to misin- 
terpretation due to these ebbs and flows of odonate life. 

Still another factor is involved which makes general con- 
clusions as to the character of the habitat of any species of 
dragonfly dangerous: the great adaptability shown by certain 
species to thrive in very different habitats. Such differences 
are usually, if not always, associated with differences in geo- 
graphical location. For example, Libellula incesta frequents 
glacial lakes in northern Indiana, especially about growths of 
Scirpus, spatter-dock and water-lilies. It never occurs about 
small, muddy, sun-exposed buttonwood ponds; but in Ten- 
nessee it does occur about just such unattractive ponds, where 
no self-respecting Indiana mcesta would fly. A species may 
be a lake-dweller in part of its range, a pond-dweller else- 
where, and a stream-dweller at a third location. 

Another difficulty in describing habitats arises from sea- 
sonable differences or evolutionary changes in the habitats 
which may escape the casual observer. For example, a suc- 
cession of muddy pools at one season may be a deep, swift- 


Occasional Papers of the Museum of Zoology 5 


flowing stream at another season. And miles of lagoons may 
a few years before have been part of a swift-flowing river. 
In the first case a dragonfly species on the wing throughout 
the year may occupy the habitat because of conditions at a 
certain season, though other seasons bring conditions incon- 
gruous with the general preferences of the species; and in 
the second case, a species may survive at least for some time 
in the lagoon, though its preferences are for the stream from 
which the lagoon was formed. 

Some dragonflies, such as the Trameas, mentioned above, 
and there are many others, range far and wide. Others, such 
as the Enallagmas, apparently live their lives practically in 
one habitat. But that this is only apparent is obvious when 
one considers the sudden appearance of large numbers of 
Enallagmas at rare intervals at a small, isolated swamp, sur- 
rounded by forest, with no known habitats of the same species 
within fifty miles or more. The known distribution of many 
species of apparently very circumscribed habitats also bears 
this out. 

Species of Hetaerinaseare dragonflies of a very circum- 
scribed habitat, and no others known to me during imaginal 
life apparently remain closer to the location where their eggs 
are laid and where their nymphal lives are passed. As to 
how the distribution of the species of the genus has come 
about we know as little as we do of the origin of the species 
themselves. But taking this into account, I know of no other 
genus in the order where one may more safely record obser- 
vations on habitats with some certainty that such observations 
are definitely descriptive of the species observed, and not thé 
record of some mere accident in the life of the species. As 
an example of the latter kind, I might mention the discovery 
of a number of larvae of Pantala hymenaea successfully living 


6 University of Michigan 


and maturing in a cement water trough in an Indiana farmyard. 

Moreover, Hetaerinas in the tropics are on the wing appar- 
ently throughout the year, and they are not as susceptible as 
most dragonflies to meteorological conditions. They are, 
therefore, less likely to be overlooked by the collector who 
may spend only a day or two at a particular habitat. Their 
habits and bright colors render them conspicuous, so they are 


detected by even the superficial observer. 


The facts just given permit more positive statements as to 
habitats and distribution of species of Hetaerina than is pos- 


sible for the larger number of dragonflies. 


The largest number of species of Hetaerina observed on 
a continuous short stretch of any stream is four, and on such 
streams the number of individuals is generally large. Such 
an optimum habitat may be briefly described. As stated, it is 
a stream, for all Hetaerinas live about running water.* The 
stream is small, two to six feet wide, with small waterfalls, 
many ripples and some quiet pools. It is usually in forest or 
brush, with bushes overhanging the water. In the stream may 
be a few arums, growths of Cyclanthus, or other similar 
aquatics. The stream will have any part of its course alter- 
nately in sun and shadow, and the ripples generally receive 
more sun than the pools. About such a sunny ripple and the 
adjacent courses of the stream, resting on the leaves or twig 
tips of overhanging bushes or on the aquatic vegetation, and 
flitting from one perch to another, Hetaerina, so far as we 
can observe, spends its imaginal life. 

With species of some other genera in the tropics, many of 


the Hetaerinas live about smaller streams and are more shade- 


* Rarely individuals, apparently strays, are taken about ponds or 
similar pools, or drought may reduce the stream where they occur to 
a succession of pools. 


Occasional Papers of the Museum of Zoology 7 


dwelling than any species we have in the north temperate 
region. But the larger number of stream-frequenting dragon- 
flies of both the tropical and the temperate regions are equally 
attracted by ripples. It may be the sun shines more continu- 
ously there than on the pools, that food is more plentiful, that 
luxuriant aquatic growths furnish convenient perches and the 
leaves and stems suitable receptacles for dragonfly eggs, and 
that oviposition directly in the water by other species is safer. 
At any rate, about the ripples of the larger northern stream 
Hetaerinas, Enallagmas and gomphines congregate at their 
season, while the ripples of some tiny brook in the tropical 
jungle flashes with the brilliant blues of Argias and Coras and 
the red of Hetaerinas. 

Tropical streams of about the same size as those in the 
north where Hetaerinas may occur, streams thirty feet wide 
or larger, have few, if any, Hetaerina inhabitants. In fact, 
odonate life generally is rare on these larger tropical streams. 
A few species find them congenial, and some others find suit- 
able homes adjacent to the river in little habitats such as pools 
at the face of a cliff, a log jam, or a bit of old river bed, but 
not parts of the river habitat itself. These larger tropical 
streams rarely attract Hetaerinas. 

The small streams where the bulk of Hetaerina life in the 
tropics is found are probably in most cases more recent than 
the species of Hetaerina which frequent them. If this is true, 
such streams received their Hetaerina faunas by migration. 
As to how this has come about, as stated before, we know as 
little as we do about the origin of the: species themselves. 
Certainly, the species are not or have not always been as 
bound to their restricted habitats as our observations would 


indicate. 


a University of Michigan 


The distribution of Hetaerinas on a particular stream may 
be such as to still farther complicate any ideas we may have 
as to the manner of distribution. For example, suppose there 
are several small'streams flowing in the same direction from 
the same range of hills and crossing a narrow coastal plain 
to enter the sea. It is obvious that the streams will approach 
their neighbors on either side very closely at their ramified 
heads. Moreover, these headwaters will approach very closely 
the headwaters of streams flowing in the opposite direction 
from the range of hills. Given a species of Hetaerina on one 
of these streams, it is obvious that it might reach adjacent 
streams across the coastal plain or by the narrowly divided 
headwaters, which would also permit the extension of the 
species to the opposite side of the range of hills. These two 
courses offer apparently the easiest paths for the dispersal 
of the species. 

But the problem is not so simple. Let us look at the Rio 
San Esteban in Venezuela. Where it leaves the rocks to flow, 
still swiftly, in its sandy bed across the coastal plain, Hetae- 
rina caja is abundant. Upstream among the rocks and in the 
lower courses of the tributary quebradas Hetaerina macropus 
is found. And at the heads of these quebradas, where the 
rock masses are the roughest and most precipitous, lives the 
largest and handsomest of the San Esteban Hetaerinas, capi- 
talis. And on the opposite side of the mountains at Bejuma, 
for example, in the Orinoco drainage, we find the same distri- 
bution of the same species. Furthermore, at Maraquita, far 
away in the interior of Colombia, seven or eight hundred 
miles up the Magdalena River, we find the same three species 
with the same distribution. 

We may assume that these three locations, with streams 


of similar character, have been in the line of similar flows of 


Occasional Papers of the Museum of Zoology 9 


odonate life. Over other streams these flows have passed, 
leaving no trace. For example, three or four kilometers back 
from the Rio Magdalena at Puerto Berrio is a muddy forest 
stream which we found very rich in odonate life, but we 
found not a single Hetaerina there, though at Cristalina, dis- 
tant about 25 kilometers, four species occur, one of them in 
large numbers. A similar muddy stream without Hetaerinas 
was found near Cumuto in Trinidad. In British Guiana are 
species of Hetaerina which would very probably find the 
Puerto Berrio and Trinidad streams congenial, but these spe- 
cies have never flowed over the far distant similar habitats in 
Colombia and Trinidad. The Hetaerina fauna of a tropical 
stream is determined by its congeniality or lack of it for the 
various species which, because of its geographical position, 
are enabled to reach it. 

The character of a stream, its rate of flow, the temper- 
ature and composition of its water, the geology of its bed, its 
fauna and flora, are all subjects which may be studied and 
determined. But the means or methods by which its plant 
and animal inhabitants have reached it are not so readily 
analyzed. As stated before, as to the dispersal of dragonfly 
species we know as little as of their origin, and the problem 
is difficult, if not impossible, of solution. Referring again to 
the Rio San Esteban in Venezuela, we find three species of 
Hetaerinas definitely distributed on the stream. These are 
caja, macropus, and capitalis. To the west across the moun- 
tains, at the swift streams about Tachira, we find two of these, 
macropus and capitalis, and a few kilometers lower down at 
La Fria, where the streams run out into sand, we find macro- 
pus and caja. But at both Tachira and La Fria occur also 
the widely distributed miniata, which is not known east of 
the Catatumbo River basin, in which both Tachira and La 


10 University of Michigan 


Fria are located. At Cristalina, in the Magdalena basin, all 
four species, caja, macropus, capitalis and miniata, occur. 
Why should miniata have stopped at the Catatumbo when the 
apparently more specialized caja ranges far to the east and 
west ? 

Another genus of dragonflies which is as closely confined 
to streams as Hetaerina is Heteragrion. In the San Esteban 
and Yaracuy valleys and over the mountains from them in 
the Orinoco drainage occurs Heteragrion chrysops alone. In 
the Magdalena basin, far to the west, occurs Heteragrion 
nutratum and three other species, but not chrysops. But in 
the Catatumbo basin chrysops and mitratum both occur. If 
chrysops could come as far west as the Catatumbo and mitra- 
tum could come as far east, and both find a congenial habitat 
there, why has not mitratum extended its range to the east, 
over the path chrysops has come; and why has not chrysops 
ranged westward to the Magdalena by the same route muitra- 
tum has travelled? 


SPECIES AND LOCALITIES 


In a former paper (Misc. Pub., Mus. Zool., Univ. of 
Mich., No. 9g) I have listed the several collecting trips. to 
the American tropics in which I have had a part or from 
which I have obtained the dragonflies for study. This paper 
deals with the Hetaerinas, numbering over 3,500 specimens, 
which have been taken on these trips. The Guatemalan Hetae- 
rinas, collected in 1905, have been fully dealt with by Dr. 
Calvert in the B. C. A., and they are not again referred to 
in this paper. The records from a single stream in Honduras, 
collected the same year and already incorporated in the B. C. 
A., are, however, again listed for purposes of comparison. The 


large lot of material on which this paper is based has been 


¢ 


Occasional Papers of the Museum of Zoology II 


identified, labelled and arranged for distribution to students 
by Mr. J. H. Williamson. He also prepared in tabular form 
a complete list of localities, with the species taken at each 
locality. This tabulation was invaluable in the preparation of 
this paper. Dr. Calvert and Dr. Ris have checked Mr. Wil- 
liamson’s determination of species from a set of specimens 
sent to each, and the determination of three species, Jaesa, 
moribunda, and mortua, rests upon Dr. Ris’s authority. 

In this paper, when reference is made to the Hetaerinas 
of Venezuela or of Colombia, for example, only the species 
taken on the trips mentioned in the first paragraph are implied. 
It is not the purpose of this paper to discuss other hitherto 
published records either of species or localities. Neither does 
this paper deal with specific characters or relationships. ‘Too 
many species of Hetaerina are wanting in my collection to 
render such an attempt practical. Of the species discussed 
in this paper I have retained large series of specimens for my 
collection, and these are available at any time for any student 
who will undertake a systematic revision of the genus. 

The localities represented by the specimens before me may 
be conveniently summarized as follows: In Guatemala, a 
section across the country on the line of the transcontinental 
railroad; two stations\on the railroad in Panama; the Santa 
Marta region and the valley of the Magdalena to just above 
Honda in Colombia; the north coast of Venezuela west of 
La Guiara, a few nearby tributaries of the Orinoco, and sta- 
tions in the Rio Catatumbo drainage south of Lake Mara- 
caibo; streams in northern ‘Trinidad; two stations in the 
department of Junin, Peru; and stations in the Demerara and 
Essequibo river basins, all within one hundred and fifty miles 
of the coast, in British Guiana. All localities are alphabetic- 


ally arranged and briefly described in the last part of this paper. 


12 University of Michigan 


Central America has a rich Hetaerina fauna with sixteen 
known species. All five of the Colombian species listed in this 
paper occur also in Central America; the four Venezuelan 
species occur in Colombia and Central America; and the two 
species known from Trinidad also are common to Central 
America, Colombia and Venezuela. But of the three Peru- 
vian species before me, only one occurs also in the material 
from the regions mentioned in the preceding sentence; and 
the four species from British Guiana are not represented at 
all in material from the other regions. 


Awaiting a careful study of the relationships of the Hetae- 
rina species, such as Dr. Kennedy has recently made of the 
Libellula species (Entomological News, XX XIII, March and 
April, 1922), the species here discussed are arranged alpha- 
betically in the text. 

These fifteen species of MHetaerina are alphabetically 
arranged and consecutively numbered in the list below. Under 
the discussion of each species, following each locality, are 
numbers in parentheses indicating the other species of Hetae- 
rina taken in that locality. For example, under H. americana, 
Guatemala, Agua Caliente (5, 9), indicates that at Agua 
Caliente, in addition to americana, cruentata and macropus 


were taken. The species are: 


1. H. americana 9. H. macropus 

2. H. caja 10. H. miniata 

3. H. capitalis 11. H. moribunda 
4. H. charca 12. H. mortua 

5. H. cruentata 13. H. pilula 

6. H. dominula 14. H. sanguinea 

7. H. fuscoguttata 15. H. titia 

S. H. laesa 


Occasional Papers of the Museum of Zoology 13 


1. Hetaerina americana Fabricius. 
Guatemala, Agua Caliente (5, 9), Amatitlan (5), Gualan (9, 15). 


In Guatemala, as well as in North America, this species 
frequents larger streams, usually fifteen feet or more in width. 
In the Motagua River basin it does not seem to descend lower 
than Gualan, which point marks the highest extension up the 
valley of H. titia. 

2. Hetaerina caja Drury. 

Panama, Rio Mazamba (10), Rio Sardanilla (7, 9, 10) : Colombia, 
Aracataca, Bolivar (3, 9), Cristalina (3, 9, 10), Don Jaca, E) Banco, 
Fundacion, Maraquita (3, 9, 10), Rio Frio (9), Santa Marta (9), 
Sevilla; Venezuela, Bejuma (3, 9), Boqueron (9), Caserio Silva (9), 
El Guayabo (10), La Fria (9, 10), Nirgua (3, 9), Palma Sola (9), 
Salom (9), San Esteban (3, 9), San Felipe (9), Tucacas; Trinidad, 
Arima, Chaguanas, Cumuto, Cunapo River (9), Diego Martin River 
(9), Maracas River (9), San Juan River (9), St. Joseph River (9). 

This species and H. macropus, in about equal numbers, 
are each represented in our collections by many more speci- 
mens than is any other species. It seems to prefer more 
exposed streams than its associates, occurring on the sunniest 
stretches of streams where their flow is slower and the bed 
less rocky. In a hilly country such habitats are found at the 
edge of the hills where the streams pass into level valleys. 

Two fairly well-defined series are represented in the mate- 
rial; a larger, with darker thoracic markings and more exten- 
sive and darker basal wing markings in the male; and a 
smaller, with paler thorax and with reduced and paler basal 
wing markings in the male. The former occurs in Panama 
and Colombia and in the Catatumbo region (La Fria and EI 
Guayabo) in Venezuela; the latter occurs in Venezuela east 
of the Catatumbo region and in Trinidad. Our opinion that 
only a single species is represented in the lot has been con- 
firmed by Dr. Calvert and Dr. Ris. The red spot on the apex 


14 University of Michigan 


of each hind wing varies in specimens from the same locality 
from one-half the usual size to the farger normal-sized spot. 
A male from Santa Marta has one front wing with a small 
apical red spot, and another male from the same locality has 
both front wings so spotted. 
3. Hetaerina capitalis Selys. 


Guatemala, El Fiscal (5, 9) ; Honduras, San Pedro Sula (5, 9, 10) ; 
Colombia, Bolivar (2, 9), Cincinnati (5, 9), Cristalina (2, 9, 10), 
Maraquita (2, 9, 10); Venezuela, Aroa (9), Bejuma (2, 9), Nirgua 
(2, 9), San Esteban (2, 9), Tachira (9, 10).° 


The Central American specimens have the thoracic dark 
markings more metallic green than the others where the mark- 
ings in the males are usually a reddish or purplish black, and 
the females are sometimes similarly colored. There is con- 
siderable 'variation in size, but this is independent of locality. 
Two males from Cristalina measure, respectively, abdomen, 
34 and 41, and hind wing, 26 and 30. 

At San Esteban we noted: “Flies sometimes in dark places 
and alights near the water on rocks; under these conditions 
very hard to see or to follow flight.” Capitalis occurs in the 
darkest habitats in which we have found Hetaerinas in the 
tropics. Such habitats frequently occur at the extreme head- 
waters of rocky quebradas. It is not impossible that our fail- 
ure to find capitalis at certain localities where its invariable 
associate, macropus, flies, as, for example, in Trinidad, may 
have resulted from inadequate exploration of the streams to 
their headwaters. 

4. Hetaerina charca Calvert. 

Peru, Colonia del Perené (9, 14). 

From the limited data available charca, like capitalis, seems 
to prefer the headwaters of the quebradas on which it occurs. 


— 


5. Hetaerina cruentata Rambur. 


Guatemala, Agua Caliente (1, 9), Amatitlan (1), El Fiscal (3, 9); 
Honduras, San Pedro Sula (3, 9, 10) ; Colombia, Cincinnati (3, Q). 


Occasional Papers of the Museum of Zoology 15 


This species occurs in the Cauca Valley, and I am at a loss 
to explain its absence ‘at Cristalina, where caja, capitalis, 
macropus, and miniata occur, when the stream at San Pedro 
Sula, almost identical in character with some of the Cristalina 
streams, has the same identical Hetaerina fauna, except that 
cruentata replaces caja. ‘This is especially puzzling since 
cruentata seems a more adaptable species than caja. 

The Santa Marta (Cincinnati) specimens are distinctly 
larger and have darker thoraces than any of the other speci- 
mens. The altitude at which we found this species at Cin- 
cinnati and our failure to find it nearby at lower elevations, 
though it is found at a much lower elevation to the north in 
Honduras, indicates that in the Santa Marta Mountains an 
isolated colony of the species is working out its destiny along 
new lines. The Santa Marta males vary from abdomen, 
39-43, and hind wing, 30-34, but we found only one this small, 
as the average size is abdomen about 42 and hind wing about 
33, about the size, by the way, of the Colombian specimens 
Hagen called lineata. Bogota and Cauca specimens do not 
differ from Central American specimens, and among these the 
males measure, abdomen about 35 and hind wing about 206. 
6. Hetaerina dominula Hagen. 

British Guiana, Rockstone (8, 11), Tumatumari (8, 11, 12), Wis- 
mar (II, 12). 

In some highly colored males, in which the basal red area 
of the wings is strongly developed, the front wings, as well 
as the hind wings, have a well-defined, distinct apical red spot, 
but this is smaller than the corresponding spot in the hind 
wings. Males evidently slightly teneral have the apex of the 
_front wings clear and the apex of the hind wings without a 
trace of red, but slightly brown-clouded. In some adult males 
the red tip of the hind wings is narrowly black or dark-edged 
apically. 


16 University of Michigan 


7. Hetaerina fuscoguttata Selys. 
Canal Zone, Panama, Rio Sardanilla (7, 9, 10). 
8. Hetaerina laesa Hagen. 
British Guiana, Rockstone (6, 11), Tumatumari (6, 11, 12). 


Dr. Ris writes that he has this species from Para, Prata 
(100 kilometers east of Para), Obidos, and a large series 
from the Tapajos. “The Tapajos series is different in that 
only a single male has the wing bases red as it is in all the 
other specimens; in these Tapajos males the wing bases are 
yellowish, though the specimens are apparently mature. I had 
long ago identified the specimen from Para as laesa, and I 
believe this determination is correct.”—Ris. In the Mon. 
Calop. the front wings are described as minutely tipped red. 
This red is not evident in the British Guiana specimens. 
9g. Hetaerina macropus Selys. 


Guatemala, Agua Caliente (1, 5), El Fiscal (3, 5), Gualan (1, 15), 
Los Amates (10, 15), Santo Tomas (10, 13, 15) ; Honduras, San Pedro 
Sula (3, 5, 10); Canal Zone, Panama, Rio Sardanilla (2, 7, 10); 
Colombia, Bolivar (2, 3), Cincinnati (3, 5), Cisneros, Cristalina (2, 3, 
10), Maraquita (2, 3, 10), Rio Frio (2), Santa Marta (2); Venezuela 
Aroa (3), Bejuma (2, 3), Boqueron (2), Caserio Silva (2), La Fria 
(2, 10), Macuto, Nirgua (2, 3), Palma Sola (2), Salom (2), San 
Esteban (2, 3), San Felipe (2), Tachira (3, 10); Trinidad, Cunapo 
River (2), Diego Martin River (2), Maracas River (2), San Juan 
River (2), St. Ann River, St. Joseph River (2); Peru, Colonia del 
Perené (4, 14). 


This is a widely distributed and apparently very adaptable 
species. It is also remarkable in the great variation in the 
development of the stigma. As in the lot of H. caja, but 
not quite so distinctly, there are two series in the present 
material of H. macropus, one series having the basal wing _ 
markings of the males darker and more extensive, and the 
other with this color more restricted and paler. The first 
series occurs in Central America, Colombia, and the Cata- 


Occasional Papers of the Museum of Zooloyy 17 


tumbo region of Venezuela. The second series occurs east 
of the Catatumbo region in Venezuela and in Trinidad. In 
their distribution the two series thus correspond to the two 
series of H. caja. This paler fauna east of the eastern Cor- 
dillera is probably correlated with a drier atmosphere than 
that of the large river valleys to the west. 

_ The three males from Peru are still more distinct and 
indicate a departure from the more typical form along the 
same lines as those of the Santa Marta Mountains form of 
H. cruentata, These specimens are the largest and richest 
colored of the entire lot. The character of size is, however, 
not as striking as in the case of the specimens of H. cruen- 
tata referred to above. Specimens of H. macropus of differ- 
ent sizes occur in each locality where the species is found, 


and this variation, except in the case of the Peru specimens, 
seems independent of locality. Specimens from Panama and 


Trinidad, for example, measure the same (male, abdomen 36, 
hind wing 25 or 26), and specimens from the Santa Marta 
Mountains (Cincinnati), Maraquita and Caserio Silva vary 
from that size up to males with abdomens 4o or 41 and hind 
wings 27 or 28. The Peru males, however, have the abdomen 
43 and the hind wing 30, a size not attained by any other 
specimens before me. 

It has hitherto been difficult or impossible to separate the 
females of this species from certain females of H. titia. In 
the female of macropus, at about midlength of the middle 
lobe of the prothorax, on ‘either side just above the suture 
between the pronotum and the propleuron (that is, about on 
the level of the lateral extremities of the front and hind lobes 
of the prothorax), there is a small but distinct outwardly 
directed antenna-like projection. At the same place on the 
prothorax of the female of H. titia there is a scarcely discern- 


18 University of Michigan 


ible elevation or small knob. When the species of Hetaerina 
are carefully studied, other valuable specific characters will 
be found in the prothorax. 

In macropus the brown tips of the wings. are more evident 
in teneral specimens than are the red tips in tenerals of other 
species where the hind wings or all four wings are red-tipped 
in the adults. 

10. Hetaerina miniata Selys. 


Guatemala, Los Amates (9, 15), Puerto Barrios (15), Santo Tomas 
(9, 13, 15) ; Honduras, San Pedro Sula (3, 5, 9) ; Canal Zone, Panama, 
Rio Mazamba (2), Rio Sardanilla (2, 7, 9); Colombia, Cristalina (2, 
3, 9), Maraquita (2, 3, 9) ; Venezuela, El Guayabo (2), La Fria (2, 9), 
Tachiraa(@ one 


The South American specimens average slightly larger 
than those from Guatemala and Honduras, and the basal red 
on the wings of the males is more extensive. The Canal Zone 
material is insufficient in quality and quantity to determine 
its status. 

This species has been recorded for relatively few localities, 
and yet at such streams as at San Pedro Sula, Cristalina, and 
La Fria it seems the most successful species. For example, 
four species of Hetaerina occur at San Pedro Sula, and of 
the total number collected there over 65 per cent were miniata; 
at Cristalina, also with four species, of the total number col- 
lected over 75 per cent were miniata; and at La Fria, with 
three species, of the total number collected over 88 per cent 
were miniata. 


11. Hetaerina moribunda Hagen. 


British Guiana, Rockstone (6, 8), Tumatumari (6, 8, 12), Wismar 
(6). 12). 


Dr. Ris writes: “I have specimens of this species from 


Occasional Papers of the Museum of Zoology 19 


Obidos and Tapajos, Brazil, and I believe it is H. moribunda.” 
12. Hetaerina mortua Hagen. 


British Guiana, Tumatumari (6, 8, 11), Wismar (6, 11). 


Dr. Ris writes: “After re-examining your specimens, rep- 
resenting a species not heretofore known to me, I am almost 
certain it is H. mortua, The description of the thoracic pat- 
tern does not fully agree, but the rest of the description and 
the figure of the appendages in the Mon. Calop. seem to indi- 
cate your species. It is strangely similar to H. dominula, but 
there are rather striking differences in both the superior and 
inferior appendages.” In the field it will be difficult to dis- 
tinguish these two species (dominula and mortua), even with 
the aid of a small hand lens. In the males, dominula has the 
red apical spot of the hind wings duller, more diffuse and 
with some brown edging, and the post-occipital tubercles are 
low and rounded, scarcely discernible to the unaided eye; in 
mortua the apical red spot is bright, not diffuse, and with 
imperceptible brown, and the post-occipital tubercles are angu- 
lar and plainly discernible. 

13. Hetaerina pilula Calvert. 

Guatemala, Santo Tomas (9, I0, I5). 

Heretofore only two specimens of this species, one from 
Mexico and one from Guatemala (B. C. A.), have been known. 
14. Hetaerina sanguinea Selys. 

Peru, Colonia del Perené (4, 9), San Ramon. 

This species was found on the lower, slower and sunnier 
parts of quebradas, and, in its relation to charca, resembles the 
frequent relation of caja to macropus. Macropus was also 
taken at Colonia del Perené, but only at one place, and its 
distribution on streams, relative to charca and sanguinea, was 


not determined. 


University of Michigan 


15. Hetaerina titia Drury. 

Guatemala, Gualan (1, 9), Los Amates (9, 10), Morales, Puerto 
Barrios (10), Santo Tomas (9, 10, 13). 

In the Motagua Valley this species occurs from’ sea level 
at Puerto Barrios to Gualan, where it meets americana, which 
descends no farther down the valley. It was well established 
at Gualan, at Los Amates 84 per cent of all the Hetaerinas 
taken were this species, and at Puerto Barrios over 97 per 
cent were this species. If I have been correct in regarding 
tricolor as a synonym of titia, this species, when its occurrence 
at sea level in Guatemala is taken into account, has a surpris- 
ingly wide distribution to the north, where it reaches Illinois, 
Indiana, Ohio and Pennsylvania in the United States. Occurring 
at sea level in Guatemala, one might expect an extensive range 
south of Guatemala, but the species occurs no farther south 
than Nicaragua, but little farther than americana. In view 
of the great similarity in the geographical distribution of the 
two species (americana and titia), their habitat distribution 
in the Motagua Valley is difficult to understand. And in Indi- 
ana I am at as great a loss to explain their abundance on 
some streams and their absence from others. 

The amount of variation in the wing coloring of males 
from the same locality is surprising. The lightest colored 
male in the present material is from Gualan, and is interme- 
diate between figures I and 2, plate 3, B. C. A. But from 
the same locality there is a teneral male with all four wings 
brown, and therefore darker than the extreme case figured 
by Calvert (figure 15, loc. cit.). I cannot tell certainly from 
the present material, but it is probable that wings of teneral 
males are suffused with brown over a larger area than is occu- 


pied by the darker brown or black of their maturity. 


Occasional Papers of the Museum of Zoology 21 


ALPHABETICAL List oF LOCALITIES WHERE COLLECTIONS 
WERE MADE 


Generally, only Hetaerina habitats are described for each 
locality, so the discussion under each locality is in no way 
indicative of the presence or absence of other nearby dragonfly 
habitats. 

1. Agua Caliente, Guatemala, on the Rio Agua Caliente, 
a swift, stony headwater of the Motagua, where the railroad 
crosses it 20.2 miles below Guatemala City. Width about 
fifty to one hundred feet. Elevation about 3,200 feet. Col- 
lected June I-2, 1909. 

Along this swift and stony river were three species, Hetae- 
rina americana, cruentata, and macropus. Unfortunately, my 
notes give no data as to the distribution of the species on 
the stream. All of the species are well represented in the 
collection, macropus by about as many specimens as the other 
two, which occurred in equal numbers. 

2. Amatitlan, Guatemala. A town on the river draining 
Lake Amatitlan. Elevation, 4,212 feet. The river is clear, 
rapid and gravelly, and the only Hetaerinas seen at Amatitlan 
were collected on this stream. Collected June 7-10, 1909. 

Of the seven specimens taken here, six are Hetaerina 
americana and the remaining specimen is cruentata. 

3. Aracataca, Colombia. On the Santa Marta-Fundacion 
railroad, near the Fundacion terminus and about fifty-five 
miles from Santa Marta. Elevation probably about fifty feet. 
Collected at irrigating ditch near fruit company station for 
only a short time on January 9, 1917, and a single male of 
Hetaerina caja was taken. 

4. Arima, Trinidad. Most of our collecting in Trinidad 


was done at two small rivers reached by electric lines from 


22 University of Michigan 


Port-of-Spain and at streams crossed by the railroad from 
Port-of-Spain eastward to Sangre Grande. This railroad 
from Port-of-Spain runs nearly directly east, and not far 
south of the hills, to Arima, en route crossing several tribu- 
taries of the Caroni River. From Arima the railroad runs 
in a southerly direction for a few miles and then turns east 
again, which is its general direction to its terminus at Sangre 
Grande. From Arima to Sangre Grande it is far south of 
the range of hills which it skirts near Port-of-Spain, and the 
streams at Arima and eastward are correspondingly slower 
flowing. West of Arima within one mile are two small, grav- 
elly streams, where we collected on March 4, 1912. 

The streams at Arima, like the other streams east of Arima 
where we collected, yielded only Hetaerina caja. Three males 
and seven females were collected, indicating that we were not 
at the habitat where the species was most numerous and active. 

5. Aroa, Venezuela. A terminus of the railroad, about 86 
kilometers above Tucacas. Elevation, 700 feet. The sandy 
Aroa Valley here is dry, and tne native flora is largely 
destroyed. Many of the surounding hills have been burned 
over. At a greater distance from town are some fine rocky 
quebradas which rise high in the mountains and which dis- 
appear in the sand a short distance after their emergence from 
the hills. One of these is west of town, possibly two miles 
in an air line. It is two to ten feet wide, with waterfalls and 
deep pools and dense growths of Cyclanthus. About two miles 
west of this quebrada is another slightly larger and slower 
one. There is another quebrada at the Tichara mine, but it 


is rock-scoured, sun-exposed, and absolutely no good. Col- 
lected March 12-14, 1920. 


On March 12 we collected up the quebrada two miles west 
of town and during the day took only one species, Hetaerina 


Occasional Papers of the Museum of Zoology 23 


macropus. The following day J. H. Williamson started col- 
lecting high up the quebrada where we had left it the day 
before, and near its source he collected three males of capi- 
talis, the only specimens of the species taken at Aroa. We 
found macropus on all the streams. The surprising thing at 
Aroa was the absence of caja, common a few miles down- 
stream at Boqueron and thence to the coast at Tucacas. It 
is probable that the streams, near the hills as they are at Aroa, 
were too swift for caja. 

6. Bejuma, Venezuela, about 30 miles west of Valencia. 
Lies in a circular plain surrounded by high hills. Through 
this plain the Rio Bejuma meanders in a sandy or gravelly 
bed, shallow pools alternating with gentle ripples. Most of 
the valley is or has been under cultivation, and the native 
flora is largely gone. The stream is fifteen to thirty feet wide 
and bordered along much of its course with wild cane which 
reaches a heighth of twenty-five feet or more. In the sur- 
rounding hills are many small quebradas of the usual swift, 
rocky type, pools alternating with swift rapids and waterfalls. 
Plant life immediately adjacent to and in these quebradas is 
usually varied and luxuriant, but, on the steep hills above, the 
native forest is usually replaced by coffee and banana plant- 
ings. The commonest plant in the forest quebradas of 
Colombia and Venezuela where we have collected is a divided- 
‘leaf palm-like aquatic growing from a foot to three or four 
feet high among rocks in the stream. It may occur as a single 
plant, as small, scattered clumps, or in a continuous growth 
filling the stream bed for a hundred feet or more. On its 
leaves rest many of the dragonflies of these quebradas. From 
photographs and my description Mr. Ellsworth P. Killip, of 
the U. S. N. M., has identified this plant as Cyclanthus bipar- 
titus Poit. Where these quebradas debouch from the hills 


24 University of Michigan 


into the alley they usually meander as shallow, dirt-sided 
arroyos which shrink in volume before the Rio Bejuma is 
reached. We could not learn the elevation of Bejuma, which 
is probably near that of Caserio Silva, about 1,500 to 2,000 
feet. Like the Rio Chirqua, the Rio Bejuma_ eventually 
reaches the Orinoco. Collected February 13-18 and 24, 1920. 

Hetaerina caja was the commonest species at Bejuma and 
was collected every day. It occurred on the Rio Bejuma and 
the lower courses of the quebradas. Macropus occurred on 
the same quebradas, but higher up than caja, and fewer speci- 
mens were taken. Of capitalis only three males and one 
female were collected, and they were taken high up near the 
sources of two quebradas. These are the same three species, 
and the only species, taken also at Nirgua and San Esteban, 
Venezuela, and Bolivar, Colombia. 

7. Bolivar, Colombia. Residence of Mr. O. L. Flye, about 
five miles out from Santa Marta. Elevation about 50 feet. 
From Bolivar the cart road has been extended about five 
miles farther to La Tigrera, following up the Tamacal River 
most of the way. This is the same T'amacal we collected at 
Santa Marta, but betweén Bolivar and La Tigrera it is a 
rapid, rocky stream, largely in shade, and its few permanent 
tributaries are of the same general character. Most of our 
collecting was done on these tributaries, as odonate life was 
not abundant along most of the course of the Tamacal, where 
rough and angular rocks, destitute of aquatic vegetation and 
washed bare by the rapid waters, offered few opportunities 
for aquatic larval life. The tributaries were less scoured and 
had occasional pools with some semi-aquatic growths. The 
elevation at La Tigrera is about 300 feet. Collected December 


20-26, 1916. 


Occasional Papers of the Museum of Zoology 25 


Along the Tamacal above Bolivar Hetaerina caja and 
macropus were flying together, but neither species was abun- 
dant. Caja was not found elsewhere, excepting three males 
which were taken at a small pond not far from the Tamacal. 
Macropus, on the other hand, was numerous on some of the 
small permanent quebradas tributary to the Tamacal. Beyond 
the road bridge over the Tamacal above Bolivar are two such 
quebradas. They are three to four feet wide near their 
mouths and are rapid and rocky with heavy vegetation. Near 
the source of one of these quebradas we took a single female 
of capitalis, the only representative of the species seen at 
Bolivar. 

8. Boqueron, between kilometer posts 68 and 69 on the 
railroad above Tucacas, Venezuela. Elevation estimated as 
375 feet. Lies in a heavily wooded and flat or slightly rolling 
country. It is about thirty kilometers above Palma Sola, and 
while the flora is generally similar the greater elevation and 
the nearer approach to the mountains gives more variety to 
the topography. One result of this is that Boqueron has many 
beautiful streams of diverse character. South of the railroad, 
in a southeasterly direction, are successively the Aroa, the 
Chivacure and the Cabobo, the latter a larger stream than 
the Arca at Boqueron. Smaller tributaries of these streams 
are not numerous. They may be, at this season, little isolated 
pools of water or, more rarely, fine, clear quebradas, with low 
rapids and pools and frequent growths of Cyclanthus bipar- 
titus (see discussion under Bejuma). At kilometer post 70 a 
trail to the north leaves the railroad and, passing through forest, 
comes to the Rio Yumarito, a beautiful stream six to twelve 
feet wide, with some deep, broad pools. There are some gravelly 
rapids and occasional rock exposures where Cyclanthus grows. 
Boqueron is a delightful region, but we found it most disap- 


26 University of Michigan 


pointing for collecting. In seven days we failed to add a 
single additional species to our dragonfly list, and in this well- 
watered region we found only two species of Hetaerinas. 
Collected March 15-21, 1920. 


Hetaerina caja was the only species which we found on 
the main streams. On the Yumarito both caja and macropus 
occurred. Caja, which was wanting at Aroa, ten or fifteen 
miles away, was the commoner Hetaerina at Boqueron. At 
Palma Sola, below Boqueron, and where the streams were 
slower, caja was still more abundant relatively, macropus being 
represented in our collections by only two females collected 
there. At Caserio Silva, where the same two species, and 
these two orly, were found, among rockier conditions, caja 
was rare and macropus was abundant. 

9. Caserio Silva. A posada between Valencia and Bejuma, 
about seven miles out of Bejuma, Venezuela. Situated on a 
small, clear stream six to ten feet wide, the Rio La Mona, 
which is generally in the sun, but has a little native forest 
remaining on it. It is generally swift-flowing in a gravelly 
bed, with pools and ripples, but uo waterfalls. Near Caserio 
Silva it flows into the Rio Chirgua, a stream in a deep, pre- 
cipitous valley, of which the native flora has been largely 
destroyed. The Rio Chirgua is twenty to thirty feet wide 
and has pools six feet deep in it. The bed is rocky or grav- 
elly. Its waters eventually reach the Orinoco. Elevation not 
known, but probably about 1,500 to 2,000 feet. Collected 
February 20-23, 1920. 

As was to be expected from the generally swift character 
of the streams, caja was rare here and was taken only on 
the Chirgua. Macropus was much more abundant, occurring 
both on the Chirgua and La Mona. Our failure to get capi- 


Occasional Papers of the Museum of Zoology 27 


talis was probably due to the fact that no rocky quebrada 
was followed back to near its source. 

10. Chaguanas, Trinidad. A station on the Port-of-Spain- 
San Fernando railroad, twenty to twenty-five miles from 
Port-of-Spain, lying not far from the coast in a flat country. 
The single small stream we found was dry in most of its 
course, with no flowing water. Collected March 7, 1912, and 
only a single female of Hetaerina caja was taken. 

11. Cincinnati. Coffee estate, twenty miles from Santa 
Marta on Mt. San Lorenzo, Colombia. Elevation about 4,500 
feet. A heavily forested region. With Cincinnati as head- 
quarters, we collected down to 2,500 feet. Streams very 
numerous, about Cincinnati very rough and rocky, with some 
fine waterfalls. Smaller streams offered better collecting than 
the larger streams, and as we descended to lower levels and 
to the quieter streams at about 2,500 feet elevation we found 
odonate life still more abundant. Collected here December 
28-31, 1916, and January I, 1917. 

At Rauca Pluma Creek, just below the house at Cincin- 
nati, we found Hetaerina capitalis and cruentata common and 
in about equal numbers. This is at an elevation of about 
4,500 feet. The stream is in forest. It has high banks, is 
very rocky and rapid, with an average width of four feet and 
a heavy flow of water. Cruentata also occurred in still larger 
numbers on two small streams, fully exposed to the sun, as a 
result of clearing of the forest, on the so-called upper road 
not far from Cincinnati and at the same elevation. At Danta 
Creek, also near Cincinnati, it was rare. Equally rare on the 
same creek were capitalis and macropus. But on the Agua 
Dulce, about four miles distant, and at an elevation of about 
2,500 feet, macropus was abundant, capitalis was very rare, 
and cruentata was absent. 


28 University of Michigan 


12. Cisneros, Colombia. Terminus of the railroad from 
Puerto Berrio. Elevation about 3,500 feet. Most of the 
native flora about Cisneros is gone and the rough, rocky 
streams, ten to thirty feet wide, are largely exposed to the sun. 
Our one day’s collecting here February 10, 1917, was so unpro- 
ductive that we explored only one stream, the Rio Santa 
Getrudis, where we took a single female of MHetaerina 
macropus. 

13. Colonia del Perené, on the Rio Perené, Department of 
Junin, Peru. Elevation, 2,230 feet. At hacienda number one 
two streams join to form the Perené. One of these, the Rio 
Paucartambo, has tributaries of varied characters on the right 
bank just above the suspension bridge. One of these flows 
through scattered bushes from a boggy spot on the bank of 
the river. One kilometer above the bridge is a very small, 
rocky quebrada. Near hacienda number one is the quebrada 
Repressa, two to four feet wide, with many waterfalls. Near 
hacienda number two is a larger and slower quebrada flowing 
into the Rio Perené. Below and east of hacienda number one 
is an old river bed of the Perené, where there are large pools 
and a little running water, and where dragonflies were very 
abundant. On the road to San Juan, past hacienda number 
two, are some small quebradas. Collected June 4-22, 1920. 

Hetaerina macropus is represented by only three males 
taken on the wooded part of a quebrada on the trail from 
hacienda number two to San Juan. On the quebrada Repressa 
and on the quebrada flowing into the Rio Perené near hacienda 
number two both charca and sanguinea were taken, but at the 
old river bed only sanguinea was found; and at the very small, 
rocky quebrada one kilometer above the suspension bridge and 


at another similar quebrada above charca was found. ‘Thus 


Occasional Papers of the Museum of Zoology 29 


charca seems to take the place of capitalis, and sanguinea the 
place of macropus of similar Venezuelan habitats. 

14. Cristalina, on the railroad 28 kilometers above Puerto 
Berrio, the latter town a river port on the Magdalena 16334 
leagues above Barranquilla, Colombia. At an elevation of 
about 1,050 feet, Cristalina lies in a rolling forested country 
and abounds in beautiful small, clear, gravelly streams with 
many ripples and a very few small waterfalls. These streams 
vary from a foot or two to six to twelve feet in width and 
all flow into the Rio Diez-y-ses, a stream of varied character, 
15 to 30 feet wide. Collected here February 12-20, 1917. 

Hetaerina caja occurred here only on the Diez-y-ses and 
on the tributary quebradas for only a short distance from 
their mouths. It was a little more numerous than macropus, 
which was found at the same places. Capitalis was rare, 
being represented in the collections by about the same number 
of specimens as macropus, and was taken only near the sources 
of the quebrada Cristalina and a tributary of the quebrada 
Sabaleticus. Miniata, on the other hand, was abundant and 
is represented by more than three times as many specimens 
as are all the other three species together. It occurred far 
up the quebrada Cristalina and throughout its course, as well 
as at its mouth, where, and on the Diez-y-ses, it was associated 
with caja and macropus. On the larger part of the quebrada 
Sabaleticus, a beautiful gravelly stream six to twelve feet 
wide, in original forest, with long pools and ripples, it occurred 
in ereat numbers, without other Hetaerina competitors. It 
was also abundant and was the only Hetaerina taken on the 
quebrada La Camelia, after a short distance above the mouth. 

15. Cumuto, Trinidad. On the railroad between Arima 
and Sangre Grande. The stream here, a tributary of the 
Caroni, is in sand and gravel, clear, with low, short ripples. 


30 University, of Michigan 


See under Arima. Collected March 6, 8 and Io, 1912. 

At Arima and at the stations east of there where we col- 
lected there is but one species, Hetaerina caja. 

16. Cunapo River, near Sangre Grande, Trinidad. A slow- 
flowing stream, eight to ten feet wide, with clay bottom. A 
tributary of the Oropuche River, flowing to the east, and 
opposite the westward-flowing Caroni. Collected here Feb- 
ruary 27, 1912. 

Hetaerina caja was abundant on this stream, and asso- 
ciated with it we found a single female of macropus. 

17. Diego Martin River, Trinidad. A stream at the end 
of the Four Roads electric car line from Port-of-Spain. A 
fine stream which we collected from Blue Basin to the car 
line. Collected February 29 and March 3, 7 and Io, 1912. 

Most of our collecting on the Diego Martin was within a 
mile of the car line, where we found only Hetaerina caja, On 
March 3 we went to Blue Basin and collected down-stream 
to the car line, and it was on the upper part of the stream, 
and there only, that we found macropus. 

18. Don Jaca, Colombia. A clear, rocky stream about 10 
to 15 feet wide, widely exposed to the sun, on the railroad 
about 25 kilometers from Santa Marta. Elevation probably 
about 50 feet. Between kilometers 17 and 18 is a small stream 
in sand. Collected these two streams December 17, 1916. 

Odonate life was not abundant on either of these streams. 
On both the only Hetaerina collected was caja. 

19. El Banco, on the Magdalena and Cesar rivers about 
86 leagues above Barranquilla, Colombia. At the season we 
were there there was no flowing water near town except the 
rivers. Along the isolated pools of a wet weather stream in a 
forest strip surrounded by a dry, treeless plain we found a 
few species of dragonflies, some in large numbers, but Hetae- 


Occasional Papers of the Museum of Zoology ay 


rinas, as was to be expected, were very scarce. Collected here 
January 23, 1917, taking only a male and a female of Hetae- 
rina caja. 

20. El Fiscal, Guatemala. Still higher headwaters of the 
Motagua above Agua Caliente and 12.7 miles below Guate- 
mala City. Small, rocky streams in deep ravines. Elevation 
about 3,700 feet. Collected June 3-6, 1909. 

Along the so-called river north of town, on June 4, Hetae- 
rina capitalis (thirty-seven specimens), cruentata (fourteen 
specimens, and macropus (two specimens) were taken. Macro- 
pus was seen nowhere else. On the stream south of town along 
the government road only one species, cruentata, was taken, 
and of it only two specimens. On June 6 I collected again on 
the river north of town, but higher on the stream than on my 
previous visit. On this upper portion I failed to find macro- 
pus, taken lower down, and cruentata was five times as abun- 
dant as capitalis, while lower down capitalis had been more 
than twice as abundant as cruentata. 

21. El Guayabo, Venezuela. A station on the Gran Fer- 
rocarril del Tachira, the railroad from Encontrados to 
Tachira, and on the banks of the Rio Zulia. Elevation, 225 
feet. El Guayabo lies in a wide, flat, wooded valley, most 
of which near town is pasture or under cultivation. This val- 
ley extends northward, without interruption and with a pro- 
gressively lower elevation, till it passes almost imperceptibly 
into Lake Maracaibo. On the left bank of the Zulia opposite 
the town are sluggish tributaries with little flow except when 
they discharge the backed-up waters of the Zulia when the 
latter falls after a rise. About six kilometers above town on 
the right bank, and crossed by the railroad, is El Cana Fraile, 
six to fifteen feet wide, in forest. On April 20 it was a suc- 
cession of pools. On April 22, the Zulia having risen in the 


32 Umiersity of Michigan 


meantime, the Fraile was a mill-race-like stream, flowing away 
from the Zulia to some inland lake or swamp. Seven or eight 
miles east of town, in the forest, we found nearly dry and 
very muddy remains of other/such cafias leading away from 
the river. Collected April 20-22, 1920. 

On the Fraile, both before and after the rise of water in it, 
and on the cana in the forest east of the town we found a few 
specimens of Hetaerina caja and mimniata flying together, the 
former about five times more numerous than the latter. 

22. Fundacion, Colombia. End of railroad from Santa 
Marta. Elevation about 50 feet. Rio Fundacion here is a 
wide, shifting, sand-bottomed river. Irrigating ditches from 
the river furnished the only other running water at that sea- 
son. Above town about two miles, on the left bank of the 
river, was a large, nearly dry and very muddy creek or arroyo 
with widely-separated pools of stagnant water eight to ten 
feet wide. Here, associated with Perithemis and Acantha- 
grion, we collected 'the only Hetaerina seen at Fundacion, a 
few specimens of caja. Collected at Fundacion January 9-14, 
1917. 

23. Gualan, Guatemala. A station on the railroad 80.2 
miles above Puerto Barrios. Elevation, 420 feet. The Gualan 
River here is a clear, gravelly stream one hundred feet or 
more in width. Just above the railroad bridge is a small, 
gravelly tributary of the Gualan. A similar, smaller but more 
shaded stream is in the forest about a mile and a half below 
town. Opposite Gualan is the Rio Manuel, a tributary of the 
Motagua, from which the city derives its water supply. My 
notes are deficient, but as I recall it the Rio Manuel is fifteen 
to thirty feet wide, and is a rapid-flowing hill stream. Like 
the Gualan River, it was not rich in odonate life. Collected 


June 11-18, 1909. 


Occasional Papers of the Museum of Zoology 33 


This is the lowest elevation I have for Hetaerina americana 
in the Motagua drainage, and the highest station for ftitia. 
The former species is represented by a single male taken on 
the Gualan River. The same day twelve titias were taken 
along the nearby tributary of the Gualan, and the same small 
stream yielded forty-four specimens of mdacropus. At the 
small stream, a mile and a half below town, macropus and 
titia were again associated, again macropus being the more 
abundant in about the ratio of eight to one. Both species 
also occurred in limited numbers along the Rio Manuel or 
some of the artificial streams diverted from it. 

24. La Fria, Venezuela. A station on the railroad above 
Fl Guayabo. Elevation, 460 feet. La Fria lies at the edge 
of the hills in the valley which extends northward to Lake 
Maracaibo. The forest is heavy mixed growth, and north of 
town, where we explored it for miles, it was nearly flat, with 
occasional small and very muddy, swampy spots, but with no 
flowing water. East of town, and crossed by a spur of the 
railroad, is a sandy quebrada, eight to ten feet wide, with a 
good flow of water, which, however, disappears in the sandy 
soil a few miles north or northeast of town. The old stone 
road south of town goes back among the hills, and about two 
kilometers from town it crosses the beautiful little quebrada 
La Fria, which in its lower course, near the stone road, is a 
gently flowing stream, five to ten feet wide, of sand, gravel 
and boulders. Growing in the stream were many plants of a 
calla-like arum. About a kilometer and a half beyond the 
quebrada La Fria the road crosses the slightly larger que- 
brada Santiaquita. This quebrada, possibly a kilometer below 
the stone road, meets with another and slightly larger stream. 
These streams were very similar to the fine little streams 
about Cristalina, Colombia, except that possibly there were 


34 University of Michigan 


more rocks in the La Fria streams, especially in their upper 
courses. Collected April 12-18, 1920. 

Hetaerina caja, not known on the swifter streams a few 
miles above at Tachira, was present, but rare, at La Fria, 
where it was taken on the sandy quebrada east of town and 
on quebrada La Fria, a total of ten specimens altogether. 
Macropus was still rarer, with a total catch of three specimens 
on quebradas La Fria and Santiaquita. Muniata was the abun- 
dant Hetaerina at La Fria and was taken every day we col- 
lected there, except one day when our entire party was in the 
streamless forest north of town. The rarity of macropus was 
a great surprise, as it was abundant at Tachira, and the swifter 
streams at La Fria seemed well suited to it. Below La Fria, 
at El Guayabo, macropus had entirely disappeared, but the 
character of the streams there was such that this is what one 
would expect. 

25. Los Amates, Guatemala. A station on the railroad 
59.3 miles above Puerto Barrios. Elevation, 160 feet. Rio 
San Francisco, below town, is fifteen to twenty feet wide, 


sluggish, with generally overhanging banks and few beaches. 
Easily waded in low water stage on June 19, but wading dif- 
ficult or impossible following heavy rains that night. After 
these rains a small wet weather stream with abundant odonate 
life made its appearance in the forest on the east of the rail- 
road below the Rio San Francisco. Collected June 18-22, 1909. 

On June 19, collecting from the railroad bridge over the 
San Francisco up-stream a mile or more, I took three Hetae- 
rma macropus, two miniata, and forty-six titia. Macropus 


was also collected along the wet weather stream east of the 
railroad. 

26. Macuto, Venezuela. About a mile and a half east of 
La Guiara. The Rio Macuto at Macuto is a clear, swift, 


Occasional Papers of the Museum of Zoclogy 35 


rocky stream, about ten feet wide, flowing directly into the 
Caribbean Sea. Collected January 28-30, 1920, 

The only species taken on this fine little stream was Hetae- 
rina macropus. ‘The absence of a coastal plain explains the 
absence of caja, and the ‘absence of capitalis from our list is 
probably to be explained by our failure to reach the higher 
sources of the stream. 

27. Maracas River, near St. Joseph, Trinidad. We col- 
lected from the Maracas Fall, 340 feet high, down-stream sev- 
eral miles. The upper part of the stream is swift and rocky; 
the lower part, swift but slower, and gravelly. Collected 
March 5, 1912. 

Hetaerinas were rare here, and we took only two speci- 
mens of caja and three of macropus. This, of all the places 
we collected in Trinidad, was the most likely-looking for cap- 
italis, but though we failed to find this species in the island, 
it is not impossible it may yet be found high in the hills above 
any of the stations visited by us. 

28. Maraquita, Colombia. On the railroad above Honda. 
Elevation about 1,500 feet. Many fine streams are near town. 
Some of these are rocky and swift, with high waterfalls and 
steep, wooded banks. Others are generally swift, but with 
much sand and without waterfalls. They vary in size from 
tiny brooks to streams 30 to 40 feet wide. As usual, the 
smaller streams were richest in dragonflies. Collected here 
February 3-5, 1917. 

The Maraquita water supply comes from the San Juan 
River. The intake is just above a high waterfall. Above the 
intake the stream is three to six feet wide, very rocky, with 
some high “waterfalls. Its banks are high and steep, with 
many helioconias and large bamboos near the water, and high 
trees. Along this upper stretch of the river we found Hetae- 


36 University of Michigan 


rina macropus and capitalis common, and miniata very rare. 
Below the intake capitalis was not found, but caja appeared 
commonly there, macropus;was in smaller numbers, but still 


common, while miniata was very rare. 


On the Poquera River, south of town, we found only caja 
and mdacropus, both common and in about equal numbers. 
The Poquera at this date was five to fifteen feet wide, in a 
wider, more sun-exposed bed than the San Juan. The water 
was low when we were there, and we were told that it some- 
times became entirely dry. 

29. Morales, Guatemala. A station on the railroad 33.6 
miles above Puerto Barrios. Elevation estimated at about 100 
feet. A large, sluggish stream here was too deep and with 
banks too brushy for effective collecting, and the only Hetae- 
rinas taken were five specimens of titia. Collected May 27, 
1909. 

30. Nirgua, Venezuela. Conditions similar to Bejuma, but 
the country rougher and, adjacent to the town, more despoiled. 
The Rio Borria, at the foot of the plateau on which the town 
is built is eight to twenty feet wide, and a few miles above 
town is in such a deep, rocky gorge with high waterfalls and 
deep pools that it is impossible to follow the stream. Above 
the intake for the city water supply, where it flows through 
brush and small trees, the stream is six to twelve feet wide 
and is gravelly and not very swift. Three or four miles north 
of town is a typical hill quebrada in pastures, brush, and coffee 
and banana plantings. On the top of the hill, above running 
water, is a humid forest. A similar small quebrada is about 
four miles northeast of town on the road to Bejuma. Ele- 
vation of Nirgua not learned, probably 1,500 to 2,000 feet. 
Collected February 25-29, 1920. 


Occasional Papers of the Museum of Zoology 37 


On the Rio Borria and the two quebradas, both Hetaerina 
macropus and caja occurred, in every case macropus being 
the more abundant. High up the quebrada north of town a 


single male of capitalis was taken. 


31. Palma Sola, Venezuela. At kilometer post 37 on the 
railroad above Tucacas. Elevation, 120 feet. Lies in a nearly 
flat, heavily wooded country. The Aroa River here is a swift, 
generally shallow stream, fifty to sixty feet wide, bed largely 
sand and the banks adjacent to the stream generally covered 
with wild cane. Three or four miles above Palma Sola there 
is a right-hand tributary of the Aroa which is eight to ten feet 
wide and which, in its characters, is merely a miniature Aroa. 
In the forests about Palma Sola are the remains of several 
wet weather streams of considerable size. There are pools of 
water two to ten feet wide and three to one hundred feet long, 
generally with little or no flow of water between pools. The 
stream beds are generally sandy, though some are muddy 
where there are extensive heliconia growths. Collected March 
4-10, 1920. 

As might be expected, this region is not rich in Hetaerinas. 
At a sluggish quebrada about one kilometer north of town we 
found caja common, and on the same quebrada took two speci- 
mens of macropus, the only specimens of the latter species 
seen at Palma Sola. Caja was taken also on the Aroa, on its 
tributary of similar character, and on an almost dry quebrada 
crossed by the railroad to San Felipe about five kilometers 


out from Palma Sola. 

32. Puerto Barrios, Guatemala. Near sea level. Small, 
sluggish and brackish streams are reached by following the 
railroad track back from the coast. One stream, the first one 


above the roundhouse, was fresh (May 28) about half a mile 


38 University of Michigan 


above the railroad, but brackish below that point. Collected 
May 25-30, 1909. 

At this small stream Hetaerina titia was very common, 
more frequently over fresh water, but observed also over 
nearby brackish water. On a stream farther up the railroad 
track, which was entirely brackish where we explored it, no 
Hetaerinas were seen. On the stream where titia was com- 
mon two specimens of miniata were taken. No others were 
seen, 

33. Rio Frio, Colombia. On the railroad about 48 kilo- 
meters from Santa Marta. Elevation probably about fifty 
feet. A fine, clear, swift stream, the Rio Frio, 40 to 60 feet 
wide, crosses the track here. At this season it could be waded. 
Trees grew to the water’s edge. Back of Esperanza Farm is 
the quebrada de Calabacito, which disappeared in the low- 
lands, but back in the hills was a stony stream of running 
water with pools six feet wide and ten to twelve feet long, 
with about two feet as a maximum depth. Streams were gen- 
erally absent at this season in the semi-arid chaparral, but 
irrigating ditches to a certain extent took their place. Col- 
lected January 6-8, 1917. 

On the quebrada de Calabacito we found both Hetaerina 
macropus and caja, the former twice as numerous as the latter, 
but neither abundant. On the Rio Frio also both species 
occurred rarely, but here caja was more numerous than macro- 
pus. On a large irrigating ditch we found only caja, which 
flew there in large numbers. 

34. Rio Mazamba, and 

35. Rio Sardanilla, two small streams, crossed by the rail- 
road. Canal Zone, Panama. The Rio Mazamba was collected 
December 6, 1916, only in its lower part near the railroad; 
but the Rio Sardanilla was followed on December 5, 1916, 


Occasional Papers of the Museum of Zoology 39 


far back into the hills, where it is a beautiful little stream 
flowing in a rocky and gravelly bed, with pools and waterfalls. 

On the Rio Mazamba, where we spent only a short time, 
and that only on its lower courses near the railroad, we took 
only Hetaerina caja and miniata, the former common, the lat- 
ter represented by a single specimen. On the Rio Sardanilla, 
where we collected more carefully, we found a Hetaerina 
fauna very rich in species, if not in individuals. On not to 
exceed two or three miles of its course we took nine Hetaerina 
caja, five fuscoguttata, eleven macropus, and one miniata. 
Unfortunately, we failed to notice if there was any particular 
distribution of these species on the stream. 

36. Rockstone, on the Essequibo River and on the Wis- 
mar-Rockstone railroad, British Guiana. Elevation not noted. 
Just east of the hotel is a large log-jammed creek, fifteen to 
twenty feet wide in low water. A short distance below the 
railroad station is a small, muddy creek, nearly dry, a succes- 
sion of stagnant pools with no running water. On the large 
island in the Essequibo opposite Rockstone is a similar but 
drier muddy creek bed. Collected February 1, 2 and 14, 1912. 

Hetaerina doninula was taken on the large creek just east 
of the hotel and nowhere else. Only fourteen specimens were 
taken. A few lJaesa were collected on the small creek below 
the railroad station, and a larger number on the island, where 
also the single specimen of moribunda taken at Rockstone was 
found. MHetaerinas were rather rare at Rockstone, our total 
catch numbering only thirty-six specimens. 

37. Salom, Venezuela. A town on the road from Bejuma 
to Nirgua. Conditions similar to those at Bejuma. Collected 
the small, sandy and gravelly river near town for a few hours 
on February 25, 1920. The water of this stream eventually 
finds its way to the Orinoco. 


40 University of Michigan 


The to-be-expected Hetaerina caja and macropus, three 
specimens of each, were collected here. 

38. San Esteban, Venezuela. A village on the Rio San 
Esteban about six miles back of Puerto Cabello. The Rio 
San Esteban is a clear, swift, rocky mountain stream, except 
near its mouth below San Esteban, where it flows for several 
miles through a nearly level sand plain. Above San Esteban 
the valley is narrow, the bed of the stream is rock or coarse 
gravel, and there are many waterfalls. Below San Esteban 
the stream is largely in the sun. Above San Esteban almost 
the entire valley is wooded, with much coffee and cacao on 
the main stream and more native forest on the higher que- 
bradas. Many tributaries (quebradas) are encountered as one 
ascends the Rio San Esteban, and these are even rougher and 
more precipitous than the main stream. The length of the 
quebradas we explored between San Esteban and Las Quiggas, 
a village on the Rio San Esteban above San Esteban, from 
their sources in the hills to their mouths in the main stream, 
varied from less than a mile to possibly four or five miles. 
We could not obtain elevations at San Esteban, but our col- 
lections were made from nearly sea level up to possibly 2,000 
or 3,000 feet at the heads of the highest quebradas. Collected 
here February 1-9, 1920. 

Below the village of San Esteban we followed down the 
Rio San Esteban for possibly three miles. Throughout this 
course the stream is largely in cacao plantings. Near the vil- 
lage Hetaerina caja and macropus were associated, but farther 
down macropus disappeared and we found only caja. High 
up on a quebrada on the left bank of the Rio San Esteban, 
just above the intake dam above the village of San Esteban, 
we found capitalis. It was most numerous near the source of 


the quebrada, and as we came down-stream it was rarer and 


Occasional Papers of the Museum of Zoology 41 


was associated with macropus. It disappeared about a mile 
below where we first found it. We found it again under sim- 
ilar circumstances on a right-hand tributary quebrada just 
above Las Quiggas. Macropus occupied the streams between 
the habitats of capitalis and caja, overlapping where their hab- 
itats came together. It was the most abundant and widely 
distributed of the San Esteban Hetaerinas. On the rocky 
quebrada on the left bank of the Rio San Esteban, opposite 
the old Salom home, we were surprised to find no Hetaerinas. 

39. San Felipe, Venezuela. Elevation, 745 feet. The town 
lies in the broad, alluvial plain of the Yaracuy River. All the 
stream beds about San Felipe were dry except the nearby 
mountain quebradas and a few small, short spring streams. 
The quebrada from which the city derives its water supply is 
open, rocky, swift and bed-scoured. We found it impossible 
to follow the stream after a short distance above the intake 
because of the precipitous sides and deep pools. Collected 
March 2 and 3, 1920. 

Only a single specimen of Hetaerina caja was taken here, 
but macropus was common on all the streams. 

40. San Juan River, near San Juan, Trinidad. A gravelly 
stream with some boulders, swift-flowing. Collected March 
f,- LO12: 

Hetaerina |caja and macropus were both common on this 
stream, the former twice as numerous as the latter. 

41. San Pedro Sula, Honduras. Estimated elevation about 
250 feet. The city lies in a broad valley, with the nearest 
hills about two miles south of town, where a small woodland 
ravine stream flows from the hills into the valley. Along this 
small stream occurred the richest Hetaerina fauna I have seen. 


Collected February 26-28, 1905. 


42 University of Michigan 


I noted of Hetaerina cruentata that it occurred along the 
stream where it emerged from the hills, but whether or not it 
also ranged back into the hills my notes do not show. It was 
back in the hills that I found macropus, miniata and capitalis, 
but again my notes fail to show if there was any local distri- 
bution of these species on the stream. The following num- 
bers of specimens of each species were taken and will indicate 
in a general way their relative abundance: cruentata 46, macro- 
pus 10, mimata 123, capitalis 9. 

42. San Ramon, Department of Junin, Peru. Elevation, 
2,800 feet. One kilometer from San Ramon, across the Rio 
Chanchamayo, is a small, sandy-bottomed creek one to two 
feet wide, flowing through cleared land and banana plantings. 
Near San Ramon is the small quebrada Apurimae, flowing 
through open country in corn and bananas, with bushes on 
the creek’s bank. The creek bed is mud, sand and boulders. 
Collected July 12-15, 1920. 

Hetaerina sanguinea was the only species observed on these 
streams. 

43. Santa Marta, Colombia. On the coast. The Manza- 
nares River and its tributary, the Tamacal, were here largely 
exposed to the sun and flowing generally over beds of fine 
sand. Collected here December 13-19, 1916, and January 3, 
IQI7. 

Three specimens of Hetaerina macropus were taken along 
small streams in a woods along the railroad track and adjacent 
to the Tamacal. Everywhere else the only species found was 
caja, which was common, 

44. Santo Tomds, Guatemala. On the coast. Near town 
a large spring forms a pool of water at the foot of a hill. 
From this spring a short stream flows into the nearby gulf. 
Collected May 29, 1900. 


Occasional Papers of the Museum of Zoology 43 


On this small stream I collected one specimen each of 
Hetaerina macropus and miniata, three of pilula, and six of 
titia. Larger series might have been collected, but I spent 
most of my time at the pool collecting dragonflies of other 
genera. 

45. Sevilla, Colombia. A station on the railroad between 
- Santa Marta and Fundacion. Elevation probably about 50 
feet. Collected here only about an hour along an irrigating 
ditch, on December 15, 1916, and took five specimens of 
Hetaerina caja. 

46. St. Ann River, Port-of-Spain, Trinidad, reached by 
the St. Ann street car line. Above the car line about a mile 
the stream is very rocky and swift, with only a few species 
of dragonflies. Collected March 1, 1912. 

Hetaerina macropus was more abundant on this stream 
than on any other stream we collected in Trinidad, and it is 
the only stream, where Hetaerinas occurred at all, on which 
we failed to find caja. 

47. St. Joseph River, near St. Joseph, Trinidad, on the 
railroad between Port-of-Spain and Arima. Stream similar 
to the San Juan, eight to twelve feet wide, swift, gravel and 
rounded stones, with some long, swift ripples. Collected Feb- 
-ruary 28 and March 11, 1912. 

Hetaerina caja and macropus were common here, the for- 
mer about five times more numerous than the latter. 

48. Tachira, Venezuela. Terminus of the railroad from 
Encontrados on the Catatumbo River. Elevation about 1,200 
feet. Tachira lies on the mountains back of La Fria where 
the hills first begin. It is in a heavily wooded region of steep 
or precipitous mountain sides with many streams. In the 
deeper valleys are swift streams fifteen to thirty feet wide, 
with pools and many rapids, but no waterfalls. The beds of 


44 University of Michigan 


such streams are wide and exposed, and odonate life is rare 
on them. ‘Two of these streams are the Rio Lobaterita, on 
the west side of town, in a deep valley, and the Rio Uraca, 
which the railroad crosses about a mile below town. Tribu- 
tary to these streams are small quebradas of various charac- 
ters. Some are miry, sluggish streams with only a small flow 
of water. Other quebradas are rocky, with waterfalls six to 
ten feet high. More rarely there are tributaries of an inter- 
mediate character, where the fall is about three to fifteen feet 
in the hundred, with frequent little waterfalls. In such a que- 
brada the stream bed is usually small, rounded boulders, with 
some gravel. Collected April 4-11, 1920. 

Along the Rio Lobaterita were frequent diversions from 
the main stream, forming little streams which soon again 
joined the larger volume of water. At these places and about 
the mouths of small tributary quebradas Hetaerina macropus 
flew in limited numbers. On a very rocky tributary, three to 
eight feet wide, with only a small flow of water, but many 
pools and waterfalls, macropus was associated with capitalis, 
both rather rare, and macropus about twice as abundant as 
capitalis. Miniata was not seen on this stream. On a some- 
what similar but flatter quebrada with more vegetation, which 
flowed through town, and which we collected from the road 
bridge far toward its source, capitalis, macropus and muniata 
were present and numerous and occurred in about equal num- 
bers. On quebradas which the railroad crossed just below 
town macropus and miniata were common in about equal num- 
bers. Along a small, short, muddy quebrada through brush 
and heliconias in the river bottom of the Rio Lobaterita, the 
three species, capitalis, macropus, and numata, occurred, but 
the first two were rare, in about equal numbers, while miniata 
was about four times as numerous as the first two together. 


Occasional Papers of the Museum of Zoology 45 


On tributary quebradas of the Rio Uraca, a beautiful stream 
above the railroad bridge, we found capitalis and miniata in 
about equal numbers, and macropus about twice as numerous 
as the other two together. These quebradas were of the inter- 
mediate type described in the last two sentences of the pre- 
ceding paragraph. 

49. Tucacas, Venezuela. On the coast. Terminus of the 
railroad, which has three inland termini, San Felipe, Aroa and 
Barquisimeto. Back from the town is a small, boggy stream 
three to six feet wide and one to three feet deep, sides steep 
or overhanging, from which the town receives its water sup- 
ply. Adjacent to the stream were literally thousands of spider 
lilies (Crinum) in bloom and there were many wet patches of 
heliconias. Dry woods and dry heliconia patches were adja- 
cent and gave evidence of flooding during the rainy season. 
The lower part of the stream is shallower, broader, and sandy 
or gravelly, with a few low ripples. Collected March 23-25, 
1920. A few Hetaerina caja were taken here. 

50. Tumatumari, on the Potaro River, about seventy-five 
miles above Rockstone, British Guiana. Elevation not noted. 
Country hilly and heavily wooded. Above town on the right 
bank is a small, sluggish stream, known as Cashew Creek, 
two to six feet wide, with a mud bed. A similar stream with 
less flow of water is on the left river bank below the’ falls in 
the river. A trail from Tumatumari leads back into the forest 
and about four or five miles from town crosses Tiger Creek, 
a sluggish stream almost too large to wade. Further up-stream 
there is a large waterfall in Tiger Creek known as Washer- 
woman Falls. The trail from Tumatumari to Tiger Creek 
crosses a few little streams from a few inches to as large as 
three feet in width. They are generally muddy and some of 


the smaller ones lose themselves in the forest. One of these 


46 University of Michigan 


streams, about three miles out from Tumatumari, was: fol- 
lowed to its mouth in Tiger Creek. Collected February 5-13, 
1912. 

Hetaerinas were rare about Tumatumari, the total capture 
amounting to only sixty-seven specimens, thirty-six of which 
were dominula, On Cashew Creek we took dominula and 
moribunda, each represented by two specimens, and mortua, 
represented by a single specimen. On the first small creek 
out from town on the Tiger Creek trail we collected four 
specimens of dominula, and on the creek on the left bank of 
the river, below the falls, we found dominula, laesa, moribunda 
and mortua, the first represented in the collection by thirty 
specimens, the next two by one specimen each, and the last, 
mortua, by twenty-six specimens. 

51. Wismar, on the Demerara River, sixty-two miles above 
Georgetown, British Guiana. Elevation not noted. Some 
small, muddy creeks tributary to the Demerara adjacent to 
town, easily waded earlier in the day, are so backed up with 
water in the late afternoon, due to the tides, that the collector 
finds them impossible to work. There is a small wooded creek 
south of town and a smaller swamp one, rising in some low 
hills, just west of town. Below town the footpath to Chris- 
tianburg crosses a muddy, log-filled creek. At Christianburg 
there is a small, muddy creek in the brush parallel to and near 
the left bank of the canal. Collected January 30 and 31 and 
February 15 and 16, 1912. 

Only two specimens of Hetaerina moribunda were taken 
at Wismar, and our notes are not clear as to exact location. 
They were taken either on the small creek south of town, or, 
less probably, on the smaller creek west of town. Dominula, 
on the other hand, was common and was taken on both the 
streams mentioned in the preceding sentence and on the stream 
along the canal at Christianburg. 


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NUMBER I31 I*EBRUARY I0, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arsor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


THE CROCODILE IN FLORIDA 
By Tuomas BARBOUR 


That curious genius, Rafinesque, one of the most surpris- 
ing and versatile of naturalists, by some hook or crook first 
learned of the existence of a crocodile in Florida. His very 
short notice is in an excessively rare number of the Kentucky 
Gazette, of which a photostat copy, made in the Library of 
Congress, is before me. There is another original copy in 
the Library at Lexington, Kentucky. I believe Dr. Stejneger 
first noticed this observation. It consists of a few brief lines 
_ only: “Our alligators have not yet been well studied by real 
Naturalists. I suspect that many species and varieties exist 
in the Southern states. The most common species is the 
Crocodilus lucius of Cuvier; there is a sharp snout alligator 
in Florida which must be his Crocodilus acutus.” (Kentucky 
Gazette (n. s.), Vol. 1, No. 29, July 18, 1822, p. 3, col. 2.) 
Rafinesque almost surely had never seen a specimen, yet he 


made an excellent surmise as to specific identity. The Florida 


2 University of Michigan 


crocodile was not heard of again until 1869, when the first 
really scientific record of the American salt-water crocodile 
(Crocodilus acutus Cuvy.), occurring within the confines of 
the United States, appeared in the Proceedings of the Boston 
Society of Natural History (1869, p. 78). This note records 
that Dr. Jeffries Wyman exhibited the head of a crocodile, 
C. acutus, obtained from the Miami River where it enters Key 
Biscayne Bay. The skull was given to Dr. Wyman by a Mr. 
William H. Hunt, a local resident, and was, he told Dr. 
Wyman, the second to be killed at that spot. The note con- 
cludes by adding that the existence of a true crocodile had 
not been previously recognized within the limits of the United 
States. This historic skull is now preserved in the Boston 
Society of Natural History, labeled by Dr. Wyman himself, 
and is No. 2,212 of the Wyman Catalogue. 

The following year (Amer. Jour. Sci. Arts, 49, 1870, p. 
105) Dr. Wyman described in more detail how he happened 
to secure the skull, and he gives a series of careful measure- 
ments. Wyman’s friends year after year, one or another, vis- 
ited Florida with him, for he was impelled annually by ill 
health to seek a milder winter climate. Mr. George Augustus 
Peabody, of Danvers, who still lives at Burleigh Farm in 
Danvers, went with him on many of these journeys, and the 
opportunities to hear from Mr. Peabody the charming remi- 
niscences of his gifted and whimsical companion will long be 
looked back upon with pleasure. In 1869, however, Wyman 
went for what I believe was his only trip to Biscayne Bay 
when he was a guest of Mr. J. Murray Forbes on board his 
yacht “The Azalea.” Miami then was a tiny settlement of half 
a dozen houses clustered about Brickell’s store, which was 
located not far from where the Royal Palm Hotel stands now. 
There was probably not a settlement in the United States that 


Occasional Papers of the Museum of Zoology 3 


had less contact with the outer world, and it is not remarkable 
that the existence of the crocodile in Florida remained known 
to only the “conchs” of the Florida Keys and to the few white 
men who, for reasons usually good and sufficient, saw fit to 
settle or move from place to place about the coast of extreme 
southern Florida, equally anxious to avoid meeting the half 
nomadic Seminoles or a better white man with a gun. 
Curiously enough, the next record for the crocodile is the 
most northerly. C. J. Maynard, who knew Wyman and had 
heard of his discovery, and who was and is a field observer of 
rarest skill, made a trip in 1872 from the St. John’s River to 
the upper end of the Indian River. Maynard wrote a short 
account of this journey for Forest and Stream (1, 1873, p. 
162). This was reprinted as Chapter 1 of Camp Life in 
Florida, which appeared in book form, published by the 
Forest and Stream Company in 1876, and was edited by 
Charles Hallock. The little volume, now of real historic inter- 
est, contains many short tales of exploring, hunting and fish- 
ing in Florida which had had an earlier appearance in the 
journal. Maynard killed a crocodile over ten feet in length 
in a creek between Lake Harney and the head of the Indian 
River. No mention is made of the water being fresh or brack- 
ish, but the Indian River is strongly saline and the locality is 
one to which it would be by no means unlikely for a crocodile 
to stray. I know of no other definite records for the Indian 
River, although I have heard rumors of stray crocodiles hav- 
ing wandered to the southern narrows near St. Lucie within 
the last thirty years. Maynard said that this was the second 
instance on record of the capture of a true crocodile in the 
United States. Curiously enough, he misspelled the name in 
the same way in both the published accounts and thereby 
added a synonym to the already somewhat complicated 


4 University of Michigan 


synonymy of the creature. It was Crocodilus acurus. 

Messrs. C. E. Jackson and W. T. Hornaday killed the 
next crocodiles on an old slide on the shores of Arch Creek, 
in Dake County, in 1875: a giant male, 15 feet 2 inches, with 
half a foot of the tail gone, and a perfect female, 10 feet 8 
inches. These were killed on successive days on the same 
slide and were well cared for. The male still ornaments the 
United States National Museum in Washington, which, I 
believe, likewise secured the skeleton of the female. Horna- 
day found another skull of a dead individual which is, beyond 
doubt, the one now in the Museum of Comparative Zoology, 
received from Ward and labeled Biscayne Bay. In the 
account of the hunt which Hornaday published in 1875 (Amer. 
Naturl., 9, p. 504) we have the first attempt to give some gen- 
eral account of habits, abundance and distribution. Hornaday 
speaks of Wyman’s having described a skull from Florida and 
called it C. acutus, but not being familiar with the variation 
within the species, Hornaday was constrained to describe his 
specimens as representing a new species, Crocodilus floridanus. 
The various specimens which are still preserved are cotypes 
of this name, no special type having been designated. More 
material now has shown that there is no diagnostic differen- 
tiation of Florida individuals and the name has neither spe- 
cific nor sub-specific value. 

Curiously enough, in later years Hornaday evidently for- 
got the Wyman notes, for in the American Natural History 
(1904, p. 320) we read: “The presence of a true crocodile 
in Florida was not discovered until 1875, when a pair of speci- 
mens of large size were collected in Arch Creek, at the head 
of Biscayne Bay, by Mr. C. E. Jackson and the writer.” Very 
probably the fact that Hornaday believed that two species 
were involved may have led him, in error, to conclude that 


Occasional Papers of the Muscum of Zoology 5 


Wyman’s skull, identified (correctly) as C. acutus, really 
came from elsewhere than Florida. For Hornaday makes it 
clear that he did not believe that both he and Wyman had the 
same species, whereas what they had were conspecific individ- 
uals of different ages. 

The closing chapter will now soon be written and before 
many years the last Florida crocodile to be recorded will claim 
historic interest in some museum equal to that now held by 
Wyman’s Miami River skull. Happily, however, the reptile 
has not wholly lacked biographers. C. B. Cory, in his Hunt- 
ing and Fishing in Florida (Boston, Estes & Lauriat, 1896, 
p. 70 et seq.), devotes a short chapter to notes on crocodiles 
and gives some fair photographs. In 1918, however, A. W. 
and Julian Dimock published their well-named Florida 
Enchantments (Outing Publ. Co., 1918, p. 89 et seq.). This 
book contains photographs which are among the finest and 
most valuable photographic records ever made in natural 
history. 

The crocodile, dwindling yearly in numbers, still may be 
found by the persistent hunter. About Ojus Creek and in 
the mangrove sloughs between Hallandale and the ocean beach 
a fair few still persist, and in February, 1920, a little croco- 
dile was found in a small brackish pool not far from Mr. 
Michael P. Grace’s garden at Palm Beach by some of his 
grandchildren. 

In February, 1919, Paul Clark, a local taxidermist at Palm 
Beach, got one which was found in Ojus Creek, floating dead 
after a hard freeze. I saw this animal after it had been badly 
mounted, and was told that it measured 14 feet 8 inches, but 
it had evidently been violently stretched. Clark subsequently 
gained some local renown by dying from the bite of a coral 


snake with which he had played. 


6 University of Michigan 


Little crocodiles when first hatched are from nine to ten 
inches long, a little longer and much more slender than newly- 
hatched alligators. 

The extension of the crocodile’s range in southwestern 
Florida is undefined, and this offers an attractive problem for 
someone who wishes a real reason to visit a region which is 
still as wild and isolated as any so near at hand. 


NUMBER 132 FEBRUARY 10, 1923, 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arpor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


WEST INDIAN INVESTIGATIONS OF 1922 
By THomAs BARBOUR 


Mr. J. L. Peters again visited the Antilles in 1922 (Feb- 
ruary-April) in the interest of the Museum of Comparative 
Zoology. He visited St. Kitts, Nevis, Anguilla and St. Eusta- 
tius. The results of his reptile-collecting are here summarized. 
His journey was specially undertaken to secure topotypes of 
Sparrman’s early described species from St. Eustatius. In 
this he was successful. Another object of the journey was 
to determine whether it might still be possible to secure 
remains of the fossil rodent Amblyrhiza, long since described 
from Anguilla. Peters found, however, that the phosphate 
bed in which the type was found was completely exhausted, 
and it is improbable that further remains are recoverable. 
Unfortunately, he was not able to visit St. Martin, owing to 
quarantine regulations, and lack of time to secure the neces- 
sary permit prevented his reaching Barbuda. Mr. Forrest, 
to whom I have often been beholden in the past, has, how- 


2 University of Michigan 


ever, sent me a small representation from that little visited 


locality. To him my very hearty thanks are due. 


Sphaerodactylus sputator Sparrman 


To secure this species, so long in doubt, was the principal 
object of Peters’ quest. He gathered an enormous series. 
This shows that the types of Sparrman now in Stockholm did, 
beyond doubt, come from this island and that the species is 
essentially as I placed it in my recent revision (Mem. M. C. Z., 
47, 1921, p. 266). It is one of the dichromatic forms, as are 
so many of the large-scaled species—and perhaps others as 
yet little known. The types are females evidently. The males 
are much smaller than the females, uniform greyish brown 
through life, or at the most with a few fine scattered dots 
usually on the head. The females are large, bulky and with 
a great variety of broken bands, blotches and spots of vary- 
ing size. 

By the kindness of my old friend and companion, Dr. 
Carlos de la Torre, rector of the University of Havana, I am 
permitted to record a very surprising observation, based upon 
field and laboratory studies carried on by Professor de la 
Torre and his correspondent, Sefior Cabrera. Lizard eggs of 
known ancestry having been secured and hatched show that 
Sphaerodactylus elegans is nothing more nor less than the 
very young of Sphaerodactylus cinereus, while extensive field 
collecting at various seasons of the year has also shown that 
during growth the individuals pass through a stage which has 
given rise to the name Sphaerodactylus intermedius, There 
is a considerable change of habitus as well as coloration dur- 
ing this course of development. ‘The new synonymy should, 


therefore, stand thus: 


Occasional Papers of the Museum of Zoology 3: 


Sphaerodactylus cinereus Wagler 


Sphaerodactylus cinereus Wagler, Syst. Amph., 1830, p. 143. 

Sphaerodactylus elegans Macleay, P. Z. 5. London, 1834, p. 12. 

Sphaerodactylus intermedius Barbour and Ramsden, Mem. M. C. 
Zee TOlO; 47, ps 2I1. 


Fullest credit is due these two investigators for this most 
enlightening observation, and we can only hope for a full 
report upon the details of their work in the future. 

I can only offer as a partial excuse for my lack of percep- 
tion in this matter the fact that year after year I visited Cuba 
during the same months. My all-year-around visits were 
during the war, when I was otherwise occupied than with 
collecting animals. I found none of the intermediate stages 
during the ten or twelve spring visits I have made to the island. 


They were to be found at other seasons. 


Anolis bimaculatus Sparrman 


This lizard from St. Eustatius was the first of this group 
of large species with smooth ventral scales to receive a name. 
Peters found it common on St. Eustatius and secured a good 
series. The dewlap in these fresh specimens is dull greyish 
white, and as in the related forms it is very feebly developed. 

I at first thought that the series from St. Kitts and Nevis 
represented distinct species. I am, however, now convinced 
that both these islands, along with St. Eustatius, are popu- 
lated by true Anolis bimaculatus, as indicated in my West 
Indian Herpetology (Mem. M. C. Z., 44, 1914, p. 279), where 
I gave reasons for following Garman. At the present time, 
however, judging from Peters’ large series, individuals from 
Nevis average very much smaller than those from the other 
islands. The following species, each represented by several 


specimens, well merit specific recognition. 


4 University of Michigan 


Anolis barbudensis, sp. nov. 


Type: M. C. Z., No. 16,167, adult male from Barbuda, 
B. W. I. W. R. Forrest, collector and donor. 

Closely related to bimaculatus, but rich brown in color, 
with many fine anastomosing white lines giving a curious ver- 
miculate appearance, dewlap brownish; upper temporal scales 
generally decidedly larger than in the St. Eustatius form and 


the median scales of the snout very much larger. 


Anolis forresti, sp. nov. 


Type: M.C. Z., No. 16,170, an adult male from Barbuda, 
B. W. I. W. R. Forrest, Health Officer of Antigua, collector 
and donor. 

This form is very closely related to A. watts: Boulenger 
of Antigua. It may be distinguished by its larger loreals, its 
occipital separated from the semicircles, usually, by only two 
rows of scales, and by its entirely uniform grey-brown colora- 


tion. The dewlap appears to be pure white. 


Anolis gingivinus Cope 

Peters secured a large series of this species which I have 
discussed somewhat elsewhere (Mem. M. C. Z., 1914, 44, 275). 
The species is poorly differentiated at best from true bimacu- 
latus, but the facies of these fresh series show that there are 
many small but constant diagnostic characters in the two 
suites of lizards collected at about the same time and similarly 
preserved under the same conditions. The differences are 
slight, usually only visible in the average, but they, neverthe- 
less, appear and seem to presage more complete speciation. 
The individuals from Nevis, St. Kitts and from St. Eustatius 
as yet are not well enough differentiated to name. It may be 


suggested that a subspecific name would be more appropriate 


Occasional Papers of the Museum of Zoology 5 


in such a case as this. If a slight degree of differentiation be 
the basis of subspecies, then one might apply trinomials which 
do have the advantage of pointing to one of the supposedly 
related types. If intergradation be the touchstone whereby 
subspecies may be determined, then insular forms automatic- 
ally demand binomial address. This seems the most generally 
convenient method to pursue with the denizens of such island 
groups as these Antilles, and, moreover, no brief is held for 
consistency now or hereafter. Absolute overlapping—. e., 
the finding of occasional identical individuals in the ranges 
of each of any two distinct forms—does not seem often to 
occur. The gradual transitions of intergradation can only 
occur where large land masses support geographic races which 


are unseparated by naturally impassible barriers of any sort. 


Iguana delicatissima Laurenti 


Peters found this iguana rare, but, nevertheless, he secured 
two specimens each on St. Eustatius and Anguilla. The hap- 
hazard distribution strongly suggests its having been carried 
about by primitive man. Indians in various localities twist 
loose the finger and toe-nails of iguanas and then stretch out 
the tendons so that their feet may be tied over their back by 
these cords. The luckless creatures are often carried about 
thus, bound and helpless, and as they live for some time with- 
out food or drink they are the most convenient meat for 
canoe journeys. ‘Thus they may have been carried through 
the Antilles, and perhaps occasionally escaped the unhappy 


fate for which they were destined. They are excellent food. 


Ameiva erythrops Cope 
Ameiva nevisana Schmidt, Proc. Linn. Soc. N. Near So es lOCO: spa. 


Schmidt described his new species from Nevis from a 


6 University of Michigan 


single faded specimen. Peters’ four fresh examples show 
that the species is not extinct on that island, as Schmidt sup- 
posed, but, nevertheless, is very rare. Peters got a fine series 
of topotypes of Cope’s erythrops from St. Eustatius. Unex- 
pectedly, they prove to be exactly the same as the Nevis indi- 
viduals. Erythrocephala from St. Kitts, according to Peters 
now almost extinct, is represented in the M. C. Z. by the fine 
series collected by Garman. This form is really doubtfully 
distinct from A. erythrops, and I am almost inclined to con- 
sider erythrops and nevisana both synonyms of erythrocephala, 
except for the fact that the latter reaches or reached an enor- 
mously greater size. The rows of ventral plates are the same 
in number. The type of coloration is similar. The supratem- 
poral scales in erythrocephala, however, are more enlarged 
over a larger area. Thus, perhaps, it is more conservative to 
consider them distinct. 


Ameiva garmani Barbour 


Peters got a fine series of this form which was previously 
known from the unique type. It is still abundant upon 
Anguilla and conspicuously distinct. 


Ameiva griswoldi Barbour 
Specimens from Barbuda collected by my kind friend, Mr. 
Forrest, Health Officer at Antigua, are indistinguishable from 
the three individuals from the latter island upon which I 


originally based this well-defined form. 


Alsophis cinereus Garman 
Peters caught two snakes on the isle of Anguilla which 
agree well with Garman’s types of this species, which came 
from both Anguilla and St. Barts, the rows of scales about 
the middle of the body being 21 in each case. 


Occasional Papers of the Museum of Zoology 7 


Alsophis rufiventris (Dumeéril and Bibron) 


Curiously enough, contrary to the conditions existing on 
most of the Antilles, snakes were not especially rare on St. 
Eustatius, and Peters secured a beautiful series of six adult 
examples which seem referable to this form. Alsophis rijers- 
maet Cope from St. Martin’s I have not yet seen, and I can- 


not speculate as to its validity. 


Bufo marinis (Linné) 


The giant toad has been introduced into Nevis, where it 
was found to have become abundant. 


8 University of Michigan 
APPENDIX 


(Translation) 


“ «Since I was first charged by Dr. C. T. Ramsden to collect exam- 
ples of all ages of the various species of “salamanquitas” to complete 
biological collections of each form, I have captured individuals of ali 
the types known in this province [Habana], but was never able to 
find young examples of S. cinereus. Desirous of solving this enigma, 
I put into a cage about twelve examples of S. elegans taken from the 
same places where S. cinereus was common and in fact living with 
that species. The first difficulty was to provide food for my captives, 
but this was done by placing ripe fruit in the cage and breeding fruit 
flies thereon (Drosophila), upon which the little lizards fed freely. 
Thus I kept my “Salamanquitas de los santos 6 de la Virgen.” So 
called, beyond doubt, because they frequent sacred pictures which, 
being generally lit with little candles or lamps, attract insects and 
hence the lizards. The lizards are often to be found abundantly 
behind these pictures, which are also often the only ones in the poorer 
houses. The lizards thus confined soon began to lose the transverse 
bars of “elegans” and to assume the ashy color of “cinereus,” but nat- 
urally they never reached the size which is attained to by ciereus in 
a wild state. I then put up hiding places and provided food in appro- 
priate places inside my house, and released specimens of elegans 
there, which grew with rapidity.’ 

“Thus, having observed all the transitions, the full credit for this 
discovery is due to the entirely modest Sefior José Cabrera, as the 
letter which I have quoted above goes to show, and as the following 
taken from a subsequent communication dated Dec. 14, 1921, sub- 
stantiates : 


“‘Esteemed Master: I add further notes which you may revise 
so that my views may be clearly placed before the Poey Natural His- 
tory Society. First you,ask me whence the eggs which I used and 
whether I was sure that they were really those of cinereus. Now I 
may only say, when you have the opportunity catch some specimens 
of Sph. cinereus in the summer, and it is almost sure that from some 
one of them you will get an egg of from 7 to 9 mm. in length and 
from it there will appear a young which has transverse bands and is 
not ashy grey. Now also when one of these Saints pictures is 
removed from the walls of an old house one often finds groups or 
colonies of large and small individuals having the two types of col- 
oration and living together. One never finds a banded individual 
large enough to contain eggs of 8 to 9 mm.’” 


“Thus, of the more than 200 individuals propagated by Cabrera 


Occasional Papers of the Museum of Zoology 9 


every single young was banded when first hatched, and thus also I 
conclude that the small differences, other than color, mentioned by 
you can be explained by the changes in proportion and other char- 
acters which take place during growth. Cabrera also reports that 
this species has the peculiar habit of a number of individuals placing 
their eggs together in the same spot. This is usually in a cavity or 
tunnel in a board which has been made by termites (comejénes). In 
one such ‘nest’ Cabrera has found no less than 25 eggs, on one occa- 
sion, and often 8 or 10. As I have said, these eggs are from 7 to 9 
mm. in long diameter and the young when first hatched are about 14 
mm. in length of body and 15 or 16 in length of tail. The egg of 
S. notatus measures about 6 mm. and the young 12 mm, body and 14 
mm. tail. 


“The young in ‘elegans’ coloration lose their cross bands in about 
two months in captivity and do not reach fully adult size until about 
five months have passed. 

“CARLOS DE LA TorRE, Sc.D., 
“Rector, Havana University.” 


I saw Cabrera’s material in Havana at de la Torre’s house, and 
there can be no doubt as to the correctness of his conclusion. It is 
a great pity that I only received these notes today (Dec. 21, 1922), 
or I would have incorporated them in the paper which was finished 
and put in type several months ago. From my own examination of 
the material in de la Torre’s hands, I am convinced that intermedius 
is in truth nothing but an intermediate transition stage between ele- 
gans and cinereus, 


I admit frankly that my own stupidity at not having suspected 
this state of affairs is almost beyond belief. I was, however, in good 
company. Again the fortuitous nature of reptile collecting is shown 
and the disadvantages under which the visitor, albeit a frequent one, 
labors. I had not found the “elegans” and “cinereus” often together 
and was completely fooled by thinking that I was collecting before 
the breeding season, hence getting adults. I was not, however. I 
never saw one of these common laying holes or “nests” which have 
never before been observed and which form one of Cabrera’s most 
interesting observations, and one not readily made unless one live in 
the same old house, well riddled with termites, for years on end— 
and keeps saints’ pictures, with their little lighted lamps, upon one’s 
walls. I envy Cabrera his opportunity, and I can picture his simple 
home, and pleased would I be to change my abode for his for the 
next few months! We owe congratulations to Cabrera for his suc- 
cess in settling this question, and to de la Torre gratitude for encour- 
aging and transmitting them—T, B. 


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NUMBER 133 FEBRUARY IO, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


RESULTS OF THE BRYANT WALKER EXPEDI- 
TIONS OF THE UNIVERSITY OF MICHIGAN 
TO COLOMBIA, 1913, AND BRITISH 
GUIANA, 1914 


THE DIPLOPODA 


By RaLepo V. CHAMBERLIN 


The diplopods secured by this expedition form a highly 
interesting collection of sixty-seven species, of which the great 
majority have not been previously described. As in the case 
of the chilopods, most of the collecting in the present group 
was done by F. M. Gaige, as indicated in the following pages 
in connection with the various records. As it seems obvious 
that our knowledge of the diplopod fauna of the regions cov- 
ered by the expedition and of the adjacent regions is still very 
incomplete, it seems best at this time not to enter into any 
general discussion of the composition and derivation of the 


fauna. The collection from Colombia, made chiefly in the 


2 University of Michigan 


vicinity of San Lorenzo, is more extensive than that from 
British Guiana, made in the Demerara River region. No spe- 
cies is common to the collections made in the two countries. 
The species secured, separately listed for Colombia and British 
Guiana, and for St. Croix, U. §. Virgin Islands, where a few 


forms were also collected, are as follows: 


CoLOMBIA 


Glomeridesmus porcellus Gervais and Goudot. 
Siphonophora graciliceps, sp. nov. 
Siphonophora pearsei, sp. nov. 
Stemmiulus major Carl. 

Stemmiulus craurus, sp. nov. 
Stemmiulus ruthveni, sp. nov. 
Stemmiulus, sp. 

Epinannolene lorensonus, sp. nov. 
Epinannolene xestus, sp. nov. 
Epinannolene arius, sp. nov. 
Epistreptus eustriatus, sp. nov. 
Spirostreptus atoporus, sp. nov. 
Orthoporus gaigei, sp. nov. 
Rhinocricus brevipes, sp. nov. 
Rhinocricus hylophilus, sp. nov. 
Rhinocricus pycnus, sp. nov. 
Rhinocricus amblus, sp. nov. 
Microspirobolus tridens, sp. nov. 
Pycnotropis colombiensis, sp. nov. 
Pycnotropis cylindroides, sp. nov. 
Polylepiscus, sp. 

Trachelodesmus angulatus, sp. nov. 
Trachelodesmus ancylophor, sp. nov. 
Dromodesmus longipes, gen. et sp. nov. 
Colombodesmus catharus, gen. et sp. nov. 
Colombodesmus lygrus, sp. nov. 
Cormodesmus hirsutellus, gen. et sp. nov. 
Alassodesmus reductus, gen. et sp. nov. 
Trichomorpha tuberculosa, sp. nov. 
Trichomorpha rugosella, sp. nov. 
Trichomorpha setosior, sp. nov. 
Trichomorpha cutyla, sp. nov. 
Trichomorpha eusema, sp. nov. 
Trichomorpha angulella, sp. nov. 
Trichomorpha paurothrix, sp. nov. 


Occasional Papers of the Museum of Zoology 


ww 


Chondrodesmus tamocolanus, sp. nov. 
Chondrodesmus cerasinopes, sp. nov. 
Chondrodesmus virgatus, sp. nov. 
Chondrodesmus virgatus frater, var. nov. 
Chondrodesmus rugosior, sp. nov. 
Arionus ulophilus, gen. et sp. nov. 
Agnurodesmus thrixophor, sp. nov. 


British GUIANA 
Glomeridesmus orphnius, sp. nov. 
Siphonophora guianana, sp. nov. 
Siphonophora corynetes, sp. nov. 
Siphonophora relicta, sp. nov. 
Siphonotus parvus, sp. nov. 
Stemmiulus drymophilus, sp. nov. 
Stemmiulus labbanus, sp. nov. 
Prostemmiulus heterops, sp. nov. 
Typhlonannolene adaptus, gen. et sp. nov. 
Nanostreptus orthacanthus, sp. nov. 
Nanostreptus astix, sp. nov. 
Orthoporus etholax, sp. nov. 
Orthoporus walkert, sp. nov. 
Orthoporus foliatus, sp. nov. 
Orthoporus, sp. 
Rhinocricus monilicornis (Porat). 
Rhyphodesmus amphelictus, sp. nov. 
Aphelidesmus guianensis, sp. nov. 
Zigwadesmus giiananus, sp. nov. 
Zigwadesmus modestus, sp. nov. 
Guianonus ectoporus, gen. et sp. nov. 
Cliodesmus cryptopygus, gen. et sp. 1iov. 


Unitep States Vircin Is_ANnps 
Rhinocricus arboreus Saussure. 
Rhinocricus monilicornits (Porat). 
Trigoniulus lombricinus (Gerstaecker). 
List OF SPECIES 
GLOMERIDES MIDAE 
Glomeridesmus porcellus Gervais and Goudot 
Ain. Soc. Ent. Fr., ser. 2, 2, p. XXVII. 
Colombia: San Lorenzo. Five specimens in forest at 4,000 
feet. July 14, 1913; F. M. Gaige. 


One specimen under leaves at 2,500 feet. July 15, 1913. 


A University of Michigan 


Glomeridesmus orphnius, sp. nov. 


Pl. 1, agshe,.2 


This species in size exceeds the known West Indian spe- 
cies, but approaches the Colombian G. porcellus (Gervais and 
Goudot). It would seem from Brolemann’s description of 
the form he identifies as porcellus that this latter is a lighter, 
chestnut species in which the head is always darker than the 
body, and in which there is a definite pattern of lighter mark- 
ings. In the present species the body is a uniform deep fus- 
cous or black, with the head pale across vertex and labrum 
and over the postantennal impressions; there is no definite 


pattern of lighter markings on collum, ete. 


The tergites have the usual series of transverse striae or 
ridges, most of which curve caudad on the lateral wings. 
Lateral wings of tergites of posterior regions of body with 
anterior angles evenly rounded, the posterior convex below 
but acutely, though but moderately, produced above, begin- 
ning with the thirteenth or fourteenth, as shown in Pl. 1, 
lig. 2. Anterior tergites with both angles rounded (PI. 1. 
Fig. 1). 

In dorsal view the collum is broader and much longer than 
the head; collum with lower end on each side narrowly 
rounded, a deep stria above the end. (PI. 1, Fig. 1. ) Width, 
2.8 mm. 

British Guiana: Labba Creek Sand Hills; July 27, 1914; 
I’. M. Gaige. One female, of which, unfortunately, the pos- 


terior end of body is missing, taken in sandy soil of forest 
floor. Holotype, M. C. Z., 5,046. 


Occasional Papers of the Museum of Zoology 5 


SIPHONOPHORIDAE 
Siphonophora graciliceps, sp. nov. 


Pie 1, dies. 3-5 


Densely fulvous, brighter at the ends. 

Head narrow, conically narrowed forward and passing 
gradually in*o the base of the beak. Beak slender, a little 
curved, passing beyond distal end of the fifth antennal article. 
(See Pl. 1, Fig. 3.) 

Collum in dorsal view with lateral edges convex, the ante- 
rior corners oblique and the anterior margin mesally incurved. 
fen 4, Pie. 3.) 

Body not keeled. Uniformly densely hairy, the hairs 
shorter than in pearset. 

Pleurites of anterior segments with cephalomesal corner 
not produced; anterior margin extended much farther for- 
ward in its ectal portion than within, the margin evenly curv- 
ing, without teeth; posterior margin convex: (See Pl. 1, Fig. 
4.) Pleurite of posterior segments as shown in PI. 1, Fig. 5. 

Number of segments, 67. 

Length, about 16 mm.; width, .65 mm. 

Colombia: Fundacion; Aug. 7, 1913; A. S. Pearse. One 
female taken with the specimens of S. pearsei. Holotype, 


ee. 2... 5,047. 


Siphonophora pearsei, sp. nov. 
Pl. 1, Figs. 6-8; Pl. 2, Figs. 9-15 
Yellow to brown, excepting the anterior seginents, which 


are typically reddish or ferruginous, most of the body occa- 


sionally being also tinged with the same color. Antennae yel- 


6 University of Michigan 


low. Head and tergites densely clothed with moderately short 
hairs of uniform length. 

The head subglobose, abruptly rounding in to the base of 
the beak. Beak moderately short, its end about on a level 
with distal end of the fourth antennal article; essentially 
straight, being only very vaguely curved; hairs along sides all 
short, but with many long hairs beneath. (PI. 1, Fig. 6; PI. 
2, Has: Q.) 

Collum viewed from above trapeziform in outline, the ante- 
rior margin mesally incurved, as usual. 

The pleurites of anterior segments with cephalomesal angle 
rounded; the outer portion of anterior margin produced for- 
ward and presenting two low, angular teeth, of which the 
mesal one is the more acute; posterior margin widely convex. 
(See Pl. 1, Fig..7.) Posterior pleurite as shown in Pl. 1, 
Fig. 8. 

Gonopods of male as shown in Pl. 2, Figs, 12-15. 
For legs of male, see Pl. 2, Figs. 10, 11. 


Number of segments, 82 to 115. 


Length, to 40 mm. ; width, to 1.8 mm. Body typically deep, 
sometimes nearly cylindrical. 

Colombia: Fundacion; Aug. 7, 1913; A. S. Pearse. Holo- 
type, M. C. Z., 5,048. Sixteen specimens collected under and 
in rotten logs in forest. “Shoot out threads of slime when 
handled, after the manner of Peripatus.” The males have a 
lower number of segments (82-92) and are in general smaller 
than the females. 

This species approaches |S. gracilicornis Carl, which is also 
a Colombian species, in form of head, pleurites, etc., but is 


readily distinguished by the details of the male gonopods. 


Occasional Papers of the Museum of Zoology 7 


Siphonophora guianana, sp. nov. 


Pl. 4, Figs. 25-27 


Color in general dark brown. 

Head subglobose, suggesting that of pearsei, but a little 
longer and not quite so abruptly narrowed to base of the beak. 
Beak short and straight. Antenna proportionately thicker than 
in pearset, gracilicorms, etc., as shown in Pl. 4, Fig. 25. 

Body broad and depressed. Pores elevated on broad tuber- 
cles, which are more pronounced and keel-like in posterior 
region. Body uniformly densely pilose, the hairs short. 

Collum in dorsal view trapeziform, rather strongly nar- 
rowed forward; the sides straight ; the anterior margin mesally 
moderately incurved. (Pl. 41, Fig. 25.) 

Pleurites of anterior segments with mesal margin weakly 
incised at middle; cephalomesal angle not produced; outer 
portion of anterior margin carried forward as usual, obtusely 
angled, the mesal part of margin convex; caudal margin con- 
vex. (PI. 4, Fig. 26.) Pleurites of posterior segments with 
outer portion of anterior margin nearly straight, the angula- 
tion small or often obsolete, the inner weakly concave, mesal 
margin weakly incurved. (PI. 4, Fig. 27.) 


Number of segments in the type, 77. 

Length, about 32 mm.; width, 2.5 mm. 

British Guiana: Forest Sand Hills; Aug. 6, 1914; F. M. 
Gaige. One female collected on ground. Holotype, M. C. 
2. 5,050. 

One female from University of Michigan Expedition Sta- 
tion 165 without further data. 

Labba Creek Sand Hills; July 27, 1914; F. M. Gaige. In 


sandy soil of forest floor. One female. 


8 University of Michigan 


Siphonophora corynetes, sp. nov. 
Pl. 2, Bigs 162gRIN3 Shics. 17-20 


General color fulvous of ferruginous cast. 

Head rather strongly narrowed forward from base, nar- 
row in front of base of antennae. Beak straight, or nearly so. 
Antennae heavy, conspicuously enlarged distad of middle. 
(Pl 2 Hae. 17. ) 

Collum of very characteristic form, the lateral margins 
short, the anterolateral corners oblique and excavated, the 
anterior margin between them strongly bowed forward and 
mesally excised. (PI. 3, Fig. 17.) 

Body not keeled, the pores not elevated, or but weakly so, 
more especially on some posterior segments, the dorsum being 
evenly convex. 

Pleurites of anterior segments as shown in Pl. 2, Fig. 16. 
Those of posterior segments as shown in Pl. 3, Fig. 18. 

Posterior gonopods of males erect, much exceeding the 
anterior pair in length; tip rather simple, spine at base of style 
abortive. (Pl. 3, Fig. 20.) Anterior gonopods as shown in 
Pl ee Fie, 580; 

Number of segments of male type, 59. 


Length, about 14 mm.; width, 1 mm. 


British Guiana: First Mourie; Aug. 26, 1914. Under fallen 
leaves in a tree clump. One male. Holotype, M. C. Z., 5,051. 


Siphonophora relicta, sp. nov. 
Pl. 3, Figs: 21-23; Plo4, Fig: 24 
Dusky brown above, with lower border of sides, all of last 
few tergites and the head a paler, fulvoferruginous color. 
Venter and legs fulvous. 


This species is separable from corynetes by the character- 


Occasional Papers of the Museum of Zoology 9 


istic form of the collum of the latter. It resembles more nearly 
S. guianana, though a notably narrower form, standing apart 
in lacking the low keel prominence or ridges of the latter spe- 
cies. The rostrum is shorter in comparison with the length of 
the legs, and the head also proportionately a little shorter. 
(See Pl. 3, Figs. 21, 22.) Collum of similar form. 

Also different from guwianana in the form of the pleurites. 
Anterior pleurites typically as shown in Pl. 3, Fig. 23. Poste- 
rior pleurites as shown in Pl. 4, Fig. 24. 

Number of segments, 62. 

Length, 19 mm.; width, 1.8 mm. 

British Guiana: Sand Hill Forest; Aug. 24, 1914; F. M. 
Gaige. One female taken in Calladium root mass. Holotype, 
eC. Z., 5,052. 

POLYZONIIDAE 


Siphonotus parvus, sp. noy. 


Pl. 4, Figs. 28, 2 


Dusky brown above anteriorly, lighter brown caudally. 

In comparison with the West Indian S. purpureus Pocock 
this is a much smaller form, having fewer segments and much 
broader relatively to its length. Head in its form, position of 
eyes, and form of antennae nearly as in that species. (See 
Pl. 4, Fig. 28.) 

The two forms appear to be distinguishable by differences 
at the caudal end of the body. In parvus the penult tergite 
completely covers the anal tergite, excepting for the tip of 
cauda, and in lateral view the middorsal line appears convex, 
bending down more of caudal border, longer relatively to the 
preceding tergite; in purpureus the cauda is considerably 


exposed and the dorsal line of penult tergite in side view is 


10 University of Michigan 


nearly straight and is smaller in comparison with the preced- 
ing tergite. (Pl. 4, Fig. 29.) 

Number of segments, 28. 

Length, about 3 mm.; width, .7 mm. 

British Guiana: Sand Hill Forest, headwaters of Hubi- 
dibu Creek; Sept. 1, 1914; F. M. Gaige. One specimen taken 
in sandy soil. Holotype, M. C. Z., 5,053. 


STEM MIULIDAK 
Stemmiulus major Carl 
Stemmatoiulus major Carl, Mém, Soc. Sci. Nat. de Neuchatel, rgr4, 
5, p. 851, Figs. 24, 26-29, 55-61. 
Colombia: San Lorenzo, 2,000 feet; July 14, 1913. One 
female among fallen leaves. 


Also at 3,000 feet ; July 16, 1913. One female under leaves. 


Stemmiulus craurus, sp. noy. 


Pl. 5, Figs. 30-34; Pl. 6, Figs. 35-36 


Body in general blackish above with a narrow middorsal 
pale line which may be in part obscure; venter and lower part 
of sides obscure fulvous to fulvo-ferruginous. Collum and 
vertex of head with a network of dark lines over a fulvous 
background. Head obscure fulvous below and somewhat 
darker between eyes and antennae. Antennae blackish and 
legs fulvous. 

Sixth joint of antennae twice, or a little more, as long as 
wide. A single large ocellus on each side. 

Gnathochilarium of male as shown in Pl. 5. Fig. 30: of 
female as in Pl. 5, Fig. 31. 

Collum angular below; three striae, on each side of which 


only the uppermost in the male is distinct in side view, the 


Occasional Papers of the Museum of Zoology IT 


others being beneath, the end of the collum being inflexed, 
the uppermost stria more widely removed from the second 
than the other two are from each other. 

Striation of segments as usual. Caudal margins of meta- 
zonites serrate in correspondence to the striae below, the ser- 
ration weak on the sides. 

Setigerous papillae of last tergite 3+3, as usual, the setae — 
moderate. 

First, second and third legs of male as shown in PI. 6, 
Bis, 35,30; and Pl, 5,.Figs:.32 and 33. 

Gonopods of male as shown in PI. 5, Fig. 34. 


Number of segments: male, 47 to 49; female paratype, 54. 


Length of female, about 25 mm.; width, 1.8 mm. Width 


of male, 1.5 mm. 


Colombia. More definite locality not known. 1913. One 


male (type) and one female. 


Also Colombia: Summit of San Lorenzo, 8,500 feet; July 
23, 1913. A male and female. Holotype, M. C. Z., 5,054. 


Stermmiulus ruthveni, sp. nov. 


Pl. 7, Figs. 46-48; Pl. 8, Figs. 49-52 


Body fulvous ventrally over most of sides and in a mid- 
dorsal line, the latter widening triangularly caudad on each 
segment; the dorsum elsewhere dusky brown, the same color 
also extending on each segment down the anterior border. 
Last tergite dusky, excepting a pale caudal border on each side. 
Collum with posterior border dark and also a dark anterior 
band just back of a narrow, pale border; the collum in general 
elsewhere pale with a network of dark lines. Head fulvous 


below level of antennae, blackish between antennae and eyes 


12 University of Michigan 
of the two sides, the vertex areolate with pale. Antennae 
blackish. Legs fulvous. 

For form of gnathochilarium of male, see Pl. 7, Fig. 46. 


Head with a single large ocellus on each side. Sixth article 
of antennae about 2.4 times longer than wide. 

Collum of the usual form, with three deep striae below on 
each side. 

Tergites of second and third segments striate only below, 
the succeeding ones becoming more and more striate until the 
striae are found entirely across the dorsum. Middorsal stria 
distinct. Caudal margin of segments serrate in correspondence 
with the striae below, the serration weak on the sides and 


absent above. 


Last tergite with the usual six setigerous papillae, the setae 
moderate, not exceeding in length the marginal setae of the 


preceding segments. 
Gonopods of male as shown in Pl. 8, Figs. 51 and 52. 
First, second and third legs of male as shown in PI. 7, 
Figs. 47 and 48, and PI. 8, Figs. 49 and 50. 
Details at tip of third legs very nearly as in craurus. 
Number of segments, male, 54 to 56. 
Length, about 28 mm.; width, 2 mm. 


Colombia: San Lorenzo, 4,500 feet; July 3, 1913; F. M. 
Gaige. Under log; one male. Holotype, M. C. Z., 5,057. 


Also, same locality, July 22, 1913; 5,000 feet. One male 


and two females, from bromeliads on trees. 


Occasional Papers of the Museum of Zoology 13 


Stemmiulus drymophilus, sp. noy. 


Pl. 6, Figs. 37-41; Pl. 7, Figs. 42-45 


This is a dark-colored species, the body in general being 
from dark brown to black ordinarily, but little lighter on the 
sides below than above, but venter paler, often orange or 
somewhat ferruginous; covered part of prozonites irregularly 
pale, fulvous, and posterior border of segments colorless, the 
prozonites showing through. A median longitudinal dorsal 
line of ferruginous color. Antennae dark. Legs pale brown 
to ferruginous. Collum and head dark, a paler fulvous area 
of triangular shape in clypeal region. 

A single large ocellus on each side. 

Gnathochilarium of male as shown in PI. 6, Fig. 37. 

Collum rounded below, the anterior angle more widely 
rounded than the posterior one, with three long striae below 
on each side, two above the margining one. 

Striae of segments arranged in general as usual, numerous 
and distinct. Setigerous tubercles of last tergite as usual. 

First, second and third legs of male as shown in PI. 6, 
Figs. 38-41, and Pl. 7, Figs. 42 and 43. Ordinary setae of 
first legs plumose. 


Gonopods of male represented in Pl. 7, Figs. 44 and 45. 
Number of segments of male, 52; of female, 50 to 54. 
Length of female, near 28 mm.; width, 2.8 mm. 


British Guiana: Labba Creek Sand Hills; July 27, 1914; 
F. M. Gaige. One female in sandy soil of forest floor. For- 
ested Sand Hills. Collected in rotten wood and in the earth. 
Aug. 14 and 17, 1914; F. M. Gaige. One male (type) and 


nine females. 


14 University of Michigan 


Sand Hill Forest; Aug. 19, 1914; F. M. Gaige. Two 
females, in rotten wood. 

Labba Creek. First Timber Landing, Clay Jungle; Aug. 
12, 1914; F. M. Gaige. One female, under a rotten log. 

Holotype, M. C. Z., 5,059. 


Stemmiulus labbanus, sp. nov. 


Pl. 8, Figs. 53-55; Pl. 9, Figs. 56-60 


Body brown to blackish along dorsum, without any pale 
median longitudinal line; paler, light brown to dilute chestnut 
over sides and below; a paler, sometimes more or less bluish, 
annulus about the caudal border of each segment. Collum 
and first tergites typically areolate with light in a dusky net- 
work. Head similarly areolate over vertex and below anten- 
nae, blackish in a cross-band between antennae. Antennae 
blackish. Legs fulvous. 

A single ocellus on a triangular dark spot on each side. 

Gnathochilarium of male as shown in Pl. 8, Fig. 53. 

Collum with lower margin convex, the anterior angle widely 
rounded and the posterior one subrectangular. A long mar- 
gining sulcus below and up the anterior margin to the level 
of the ocellus, and above this a single short one across poste- 
rior portion of plate only. 

Striation of segments of trunk in general as usual. Pos- 
terior margin of metazonites deeply serrate below, the serra- 
tion becoming minute in going dorsad and above absent. 

Last tergite with setigerous papillae 3-++-3 as usual. 

Gonopods of male as shown in Pl. 9, Figs. 59 and 60. 

First, second and third legs of male as shown in PI. 8, Figs. 
54 and 55, and Pl. 9, Figs. 56-58. 


Number of segments of male, 52 to 53; of female, 49 to 54. 


Occasional Papers of the Museum of Zoology 15 


Length of largest female, near 30 mm.; width, 3 mm. 


British Guiana: Labba Creek, First Timber Landing, Clay 
Jungle. In rotten logs. Aug. 12, 1914; F. M. Gaige. One 


female taken under fallen leaves ina tree clump. Holotype, M. 
i, Z.5 5,004. 


Stemmiulus sp. 


Colombia: San Lorenzo. At 7,600 feet on July 19, and 
at 8,000 feet on July 23, 1913. Five small females of fifty 


to fifty-two segments taken in bromeliads on the ground. 


Prostemiulus heterops, sp. noy. 


Pl. 9, Fig. 61; Pl. 10, Figs. 62 64 


Color of body dark brown, with covered region of pro- 
zonites pale and the posterior border of metazonites transpar- 
ent, so that the light color of prozonites shows through. No 
definite markings. Legs brown, the antennae darker. 

In the type there are two ocelli on the right side and three 
on the left in a subvertical series, the dorsal one largest. (See 
Pl. 10, Figs. 63 and 64.) 

Gnathochilarium as shown in Pl. 10, Fig. 62 

The lower anterior corner of the collum on each side widely 
rounded, with the posterior corner much more narrowly 
rounded. ‘lwo long true striae or sulci on each side. (See 
Pl. 10, Fig. 63.) 

Striae of following segments disposed about as usual. The 
upper striae very oblique. The finer impressed lines of sur- 
face of collum and all other segments are longitudinal, fine 
but distinct, and numerous. Serration as shown in PI. 9, 
Fig. 61. 


Setigerous tubercles of last tergite 3+3 as usual. 


16 University of Michigan 


Number of segments, 47. 
Length, about 20 mm.; width, 2 mm. 


British Guiana: Forested Sand Hills; Aug. 17, 1914; F. 
M. Gaige. One female. Holotype, M. C. Z., 5,065. 


EPINANNOLENIDAE 


Epinannolene lorenzonus, sp. nov. 


Pl. 10, Figs. 65-67 . 


The color at present is brown with pale annuli about cau- 
dai border of metazonites, but otherwise with no distinct light 
or dark markings. Legs fulvous. 

Body of nearly uniform length, excepting where a little 
constricted a few segments back of head. 

Ocelli on each side in type nineteen or twenty in four 
series; thus, 7, 6, 4, 2, or 8, 6, 4, 2. Eyes somewhat more 
than two and a half times their diameter apart. 

Collum with lower margin on each side extending a little 
below level of the second tergite, the end bent mesad below, 
rounded, with both anterior and posterior corners also rounded ; 
four principal striae on each side, with several incomplete 
finer ones. (Pl. 10, Fig. 65.) 

Sutural constriction, as usual, sharply defined. Pore in 
contact with suture. Prozonite anteriorly with several fine 
encircling striolations. 

Segments, excepting in anterior region, longitudinally stri- 
ate only below. 

The species is most readily distinguished by the form of 
the gonopods, the distal end of which is smooth, without proc- 
esses or incisions, as shown in PI. 10, Figs. 66 and 67. The 
gonopods of the type, however, had been dry and rubbed and 


ost setae are apparently lost. 


Occasional Papers of the Museum of Zoology 17 


Number of segments, 57, or near that number. 

Diameter, 2.25 mm. 

Colombia: San Lorenzo, 3,000 feet ; July 16, 1913. “Under 
leaves and stones.’ One male, which had dried from evap- 
oration of the alcohol. Holotype, M. C. Z., 5,066. 


Epinannolene xestus, sp. nov. 
Pieter, Bic. 70 

Fuscous or nearly black, shining; on each ordinary seg- 
ment a paler vertical stripe on each side just caudad of fur- 
row and metazonite paler caudally above. Head with dark 
band between eyes, below which fulvous or ferruginofulvous, 
the vertex with network of dark. Anal valves paler than body. 
Legs light ferruginous, about like the face. The paratype at 
11,000 feet is more distinctly annulate, the obscure ferruginous 
annuli encircling segments in front of caudal border. 

Eyes about once and three-fourths their greatest diameter 
apart. Ocelli deeply pigmented, mostly large, in four series, 
about 28 in number: thus, 8, 7, 7, 6. 

Collum with lower end moderately bent mesad, not extend- 
ing below level of second tergite; lower margin rounded, as 
are also both corners, the lower margin at middle a little flat- 
tened or vaguely incurved. A deep margining sulcus below 
and up the front to level of eye; above this three deep sulci 
or striae, of which the uppermost curves up anteriorly and 
terminates near same level as margining sulcus; the second 
sulcus bends up anteriorly to end against the uppermost one, 
and the first ends similarly on the second, or the second may 
extend up parallel to the first. In addition there are two or 
three short sulci across posterior border above the others and 
there may be a short impression farther cephalad. See further 
7,11, Fig. 7o. 


18 University of Michigan 


Segmental constriction distinct and complete; marked 
across dorsum with a series of impressed punctae, these 
impressions continuing and becoming coarser down the sides 
and below extending upon the metazonite on the longitudinal 
striae. Surface of segments otherwise in general smooth and 
shining. The striae extending up the sides on the most ante- 
rior segments as usual. Pore removed from suture. 

Last tergite caudally rounded, even with the valves. Anal 


valves mesally narrowly margined. 
Number of segments (female), 52 to 54. 
Length, about 30 mm.; width, 2 mm. 


Colombia: San Lorenzo, 4,500 feet; July 3, 1913. “Under 
bark of stump near creek” and “under log.” ‘Two adult 
females and one immature one. Also at 4,000 feet; July 4, 
1913; F. M. Gaige. One’ female. Holotype; M. C..Z5 5,0ay% 


Epinannolene arius, sp. nov. 


Pl yo, Fig: 68 ‘PL’ ix, Fim Ge 


Brown, the posterior portion of metazonites and anterior 
portion of prozonites paler, as usual, lower portion of sides 
paler than dorsum. Collum dark within a pale border with 
median region covered with a network of dark over a pale 
background, a few following tergites also with a similar net- 
work of dark lines. Vertex of head covered with network of 
dark lines over a pale ground; dark between eyes, becoming ~ 
paler below. Legs light brown. 

Vertigial sulcus fine, ending on a transverse sulcus joining 
inner angles of eyes. Eyes transversely elongate, acutely 
angled at mesal ends, not fully 1.7 times their greatest diam- 
eter apart. Ocelli 18 to 23 in number, arranged in three series: 


2.6.5 7,7, 4)-20ao, Cuno: 


Occasional Papers of the Museum of Zoology 1g 


Collum strongly narrowed down the sides, each lower end 
moderately inflexed, rounded, the anterior corner more widely 
convex than the posterior. Margined below and up the front 
to eyes; above margining sulcus typically five striae, of which 
the second and third from the uppermost are complete, while 
the others may be more or less interrupted in the middle region, 
giving the appearance of seven or more distinct but mostly 
incomplete striae. (See Pl. 10, Fig. 68.) 

Segments deeply constricted; smooth, excepting below, 
where striate as usual. Pore widely removed from furrow. 

Gonopods of male as shown in PI. 11, Fig. 69. 


Number of segments of male, 47; of female, 54. 
Width of female, 1.4 mm.; of male, I.2 mm. 


Colombia: San Lorenzo, 2,500 feet; July 15, 1913. “Under 


’ 


leaves and logs.” One male and one female. Holotype, M. 


= 7. 5,070: 
Typhlonannolene, gen. nov. 


Gnathochilarium as in Epinannolene. Mandibles with nine 
to ten pectinate lamellae. Eyes, none. Repugnatorial pores 
beginning on the fifth segment. Character of male gonopods 
unknown. 


Genotype, T. adaptus, sp. nov. 


Typhlonannolene adaptus, sp. nov. 


Ph 1. Figs. 73673 


General color brown, the body distinctly annulate, the ante- 
rior portion of prozonites and the caudal portion of metazo- 
nites being paler. Commonly a darker spot showing on each 
side of each segment toward ventral surface. Legs brown. 

Antennae clavate as shown in PI. 11, Fig. 73. 


Gnathochilarium of female as shown in Pl. 11, Fig. 72. 


20 University of Michigan 


Collum inflexed at ends below. Strongly narrowed ventrad, 
the ends narrow, rounded. ‘Typically with six longitudinal or 
sublongitudinal striae on each side of which the two upper- 
most are longer and more oblique than the others; striae end- 
ing in a vertical margining stria along anterior border extend- 
ing part way to middorsal line. (See Pl. 11, Fig. 71.) 

Segments in general not truly constricted, but with suture 
very distinct throughout, smooth; pore contiguous with suture, 
the latter on some a little angularly bent at its level. Meta- 
zonites striate below, the segments otherwise wholly smooth 
and shining. 

Anal tergite caudally rounded; surpassed by the valves. 
Anal valves narrowly but sharply margined. 

Number of segments, 67 to 73. 

Length, about 42 mm.; width, 2.2 mm. 

British Guiana: Second Mourie; Aug. 19, 1914; F. M. 
Gaige. ‘Two females in sand. 

Forested Sand Hills. “In rotten wood and in the earth.” 
Aug. 17, 1914; F. M. Gaige. One female. 

Also, Aug. 14, 1914. One female in sandy soil of forest 
floor. | 

Labba Creek Sand Hills ; July 27, 1914; F. M. Gaige. Tiree 
iemales collected in sandy soil of the forest floor. 

Sand Hill Forest; Aug. 22, 1914; F. M. Gaige. A female 
taxen in Calladium root mass. 


Holotype, M...C. Z.,, 5,072. 


SPIROSTREPTIDAL 
The following artificial key will aid in separating the spe- 
cies of the family described in this paper. It is based pri- 


marily upon the females. 


Occasional Papers of the Museum of Zoology 21 


a. Eyes twice or more their greatest diameter apart. 


b. Repugnatorial pores contiguous with segmental sutures. 
: S. atoporus. 


bb. Repugnatorial pores well removed from sutures. 


c. Last tergite transversely sulcate; segments sixty-five. 
N. astix. 


cc. Last tergite not transversely sulcate; segments fifty-five 
to fifty-six. 
d. Collum angled below, with four striae; anal scale with 


caudal margin only weakly convex, not mesally angled. 
N. orthacanths. 


dd. Collum with lower margin straight, only two striae; 
anal scale with caudal margin strongly angled. 
N. gracilior. 


aa. Eyes clearly less than twice their greatest diameter apart. 
b. Segments of middle and posterior region of body striate up to 


or nearly to pore; suture strongly ribbed throughout; eves 
about 1.4 times their diameter apart. E. eustriatus. 


bb. Segments of middle and posterior region of body striate only 
beneath; eyes less than their diameter apart. 
c. Posterior angle of last tergite and of anal scale obtusely 
rounded. O. gaiget. 
cc. Posterior angle of last tergite and of anal scale acute. 

d. Last tergite with caudal portion set off or crossed by 
one or two sharply impressed transverse sulci; seg- 
ments fifty-eight to sixty-one. Anterior angle of col- 
lum of female produced, 

e. Second stria curving upwards in front as usual; 
next to uppermost stria very short, crossing only 
caudal border. O. walkeri. 

ee. Second stria short, straight, not curving upwards 
anteriorly; next to uppermost stria complete, not 
there abbreviated. O. foliatus. 

dd. Last tergite at most with a very shallow transverse 
depression, no sharply impressed sulcus, segments 
fifty-four to fifty-five. Anterior angle of collum of 


female subrectangular, male conspicuously produced. 
O. etholax. 


22 University of Michigan 


Nanostreptus orthacanthus, sp. nov. 


Pl. rr, Figs. 74 765.61 12, Figs, 71-81 


I;xposed portion of segments greyish blue or darker ovet 
prozonite, with metazonite encroached upon by the blue below, 
the caudal portion of the metazonite ringed with fulvous or 
ferruginous to nearly white, the light ring embracing some- 
what more than half of the metazonite or prozonite and dark 
part of metazonite both of same dark color, often nearly black. 
A dark line along each side at level of pores and a dark mid- 
dorsal line over all or much of length. Collum and head 
typically abruptly darker than the following region of body, 
deep brown or blackish, the collum narrowly bordered with 
obscure ferruginous. Antennae and legs deep brown. 

Median sulcus of head distinct only across vertex. Cly- 
peal foveolae 2+2. Eye patch wider transversely than long, 
acutely angled at mesal end; eyes about two and one-half 
times their diameter apart ; ocelli about 26 in number, arranged 
in five series: e. g., 6, 8, 5, 2, 2. Antennae short, when laid 
back on a level with the repugnatorial pores reaching the cau- 
dal edge of collum. Cardo of mandible with lower margin in 
ectal view straight excepting a slight angle at caudal end. 
(Pi. 141, Bie: 76.) 

The lower, triangular end of the collum on each side is 
bent obliquely under, the anterior edge of the inflexed portion 
being larger and more oblique than the posterior; inflexed 
surface covered by four longitudinal sulci. The longitudinal 
edge formed by bending in of lower end of collum, long and 
straight and with a single deep margining sulcus just above it. 
In situ the antenna lies in the hollow below this edge: (PI. 
11, Figs. 74, 75.) 


Suture of segments deep and coarse throughout, marked 


Occasional Papers of the Museum of Zoology 23 


across dorsum by deep and coarse punctiform impressions ; 
widely and moderately curved opposite pore. Pores small, 
each situated just in front of caudal edge of dark bend of 
metazonite. Metazonites deeply striate beneath and half-way 
up the side to the pore, the upper striae incomplete caudally, 
only crossing or but little surpassing the dark band of 
metazonite. 

Last tergite caudally rounded, greatly exceeded by the anal 
valves. Anal valves mesally compressed and strongly elevated, 
the free edges evenly convex. (PI. 12, Fig. 80.) 

Anal scale as shown in PI. 12, Fig. 81. 

Characterized especially by gonopods of male, which are 
represented in Pl. 12, Figs. 77 and 78. 

Leg of sixth segment of male as represented in Pl. 12, 
Fig. 79. 

Number of segments, 55. 

Length of male, about 40 mm.; width, 3 mm. 


British Guiana: Near Demerara River, “Cacao Plantation 
about camp.” Rotten logs. July 16, 1914; F. M. Gaige. One 
male and three females. Also, “East trail along Demerara 
River, collected in the wet earth close to the high tide mark, 
a common form.” Aug. 8, 1914; F. M. Gaige. Ten specimens. 

Labba Creek Sand Hills; July 27, 1914. In sandy soil of 
forest floor. Two females. Holotype, M. C. Z., 5,077. 


Nanostreptus astix, sp. nov. 


Pl. 12, Figs. 82-84; Pl. 13, Fig. 85 


This closely parallels in its general structure the preceding 
species. The type appears considerably lighter in color than 
those of the latter species, lacks the lateral dark lines and has 
the middorsal line absent or but obscurely indicated on some 


24 University of Michigan 


of the segments. Segments with fulvo-ferruginous band about 
caudal border embracing more than half the length of meta- 
zonite. Covered portion of prozonite also pale. The dark 
region of prozonite including numerous light spots or areo- 
lations, particularly on sides. On some of the segments the 
light color may encroach upon the dark ring so as in part 
almost to obliterate the latter. Last tergite and anal valves 
dark, excepting a narrow caudal border on each side of the 
former. Median region of collum areolated with light. Head — 
with a dark band between eyes and extending ventrad between 
antennae ; vertex areolated with light. Legs light ferruginous. 

Sulcus across vertex as usual. Eyes more elongate trans- 
versely than in orthacanthus; ocelli 57 in number, in seven 
series, thus: 10, II, II, 10, 8, 5, 2, the two of lowest series 
very small. Lower margin of cardo of mandibles concavely 
excavated as shown in Pl. 12, Fig. 83, a deep margining sul- 
cus above it. 

Lower end of collum less abruptly bent under than in 
orthacanthus, and the lower angular part bent down more 
nearly vertically below level of second tergite where the latter 
exceeds it in orthacanthus. (See Pl. 12, Fig. 82.) 

Segmental suture differing from that in the preceding spe- 
cies in being more slightly curved opposite the pore, often 
essentially straight, and more nearly smooth, not marked by 
such deeply impressed coarse punctae. Pore at or near caudal 
edge of dark ring. 

Anal scale longer in proportion to width than in orthacan- 
thus and more obtusely angled posteriorly. (See Pl. 12, Fig. 
84, in comparison with Fig. 8.) A conspicuous difference 
is also presented by the last tergite, which is more acutely 
angled behind and presents a deep transverse furrow behind 
which the plate is depressed and more roughened. Compare 


Occasional Papers of the Museum of Zoology 25 


Pl. 13, Fig. 85, this furrow less marked and sometimes obscure 
in young specimens. 

A larger species with the number of segments 65 as 
against 55. 

Length, about 70 mm.; width, 4.5 mm. 

British Guiana: Forested Sand Hills. In rotten wood. 
Aug. 17, 1914; F. M. Gaige. Two adult and five partly grown 
females. 

Sand Hill Forest; Aug. 22 and 24, 1914. Three females 
“in Calladium root masses.” 

Labba Creek Sand Hills. July 27, 1914. Two females. 

First Mourie; July 31, 1914. In humus of tree trunk. 
Two specimens. 

Second Mourie; Aug. 19, 1914. In sand. One female. 

Sand Hill Forest; Aug. 19, 1914. Two females. 

Holotype, M. C..Z., 5,081. 


Nanostreptus gracilior, sp. nov. 


PE e,- Figs. 86; 87 


A female apparently pertaining to this genus is a smaller 
and more slender form than those of the preceding species. 
It differs decidedly in not having the collum at all truly angled 
beneath, the lower margin on each side being straight as in the 
Colombian N. inconstans Carl, the corner rounded, the ante- 
rior one lower than the posterior. Three or four stories on 
each side as shown in Pl. 13, Fig. 86. 

The lower edge of cardo of mandibles only very slightly 
incurved (Pl. 13, Fig. 86). Ocelli in type 41, in six series; 
Pius, 0, 9, 8, 7, 5, 3- 


Segmental suture deep and smooth, weakly excurved oppo- 


26 University of Michigan 


site pore. Segments, excepting anterior ones, striate only 
below, as usual. 

Last tergite evenly convex, caudally obtuse, without trans- - 
verse sulcus. Anal scale strongly angled behind (Pl. 13, 
Fig. 87). 

Number of segments, 56. 

Length, about 36 mm.; width, 2.7 mm. 

British Guiana: First Mourie; July 31, 1914; F. M. Gaige. 
In humus of tree trunk. 

Labba Creek Sand Hills; July 27, 1914; F. M. Gaige. One 
female. 


Holotype, M. C. Z., 5,088. 


Epistreptus eustriatus, sp. nov. 
Pl. 13, Figs. 88, 80 


Prozonites typically cinereous or pale bluish-grey, the meta- 
zonites ordinarily darker, more brownish, with a narrow cau- 
dal border much deeper in color, ferruginous. Posterior bor- 
der of last tergite and the elevated borders of anal valves also 
darker, ferruginous. Antennae and legs fulvous. Head with 
ferruginous background; a dark band between eyes; vertex 
with network of dark lines. 

Vertigial sulcus ending in a depression at edge of the dark 
inter-ocular band. Clypeal foveolae 2+2. Eyes elongate 
transversely, being about 1.66 times wider than high, the inner 
angle acute, 1.4 times their diameter across. Ocelli about 47 
in number, in six or seven series; é@. g., II, 10, 9, 7, 5, 3, 2. 
Antennae, when laid back parallel to median line, scarcely 
reaching caudal edge of collum. Cardo of mandible with 
caudo-ventral corner of outer face acutely extended ventrad, 


the lower margin concave. 


Occasional Papers of the Museum of Zoology 27 


Collum with lateral lobe on each side not at all inflexed, 
subvertical, its anterior portion in part concealing the base of 
cardo of mandible. Lower margin weakly convex, both ante- 
rior and posterior corners rounded, the anterior one more 
widely than the posterior. On each side three striae in addi- 
tional marginal one which curves below the edge inferiorly, 
the uppermost of these striae deepest as usual. See further 
Pie13, big. 88. 

In a typical segment of the middle region of the body the 
suture is deep and wide, straight throughout or but weakly 
excurved opposite the pore, and is crossed by longitudinal 
ridges or ribs dividing it into a series of pits. Below the pore 
most of these ribs continue across the metazonite as the lower 
limiting ridges of the striae, which extend up to or nearly tc 
the pore; shorter striae or simple sutural ribs in part alternate 
with the principal striae and their limiting ridges. Pore nearly 
one-fourth the distance from suture to caudal margin of meta- 
zonite. Anterior region of prozonite with numerous regular 
cross-striolations which anastamose at intervals, excepting the 
most caudal one, or in part also the next to last, the striola- 
tions finely beaded. Segment otherwise dorsally wholly 
smooth excepting vague fine lines and punctae visible under 
good magnification. Sternites nearly smooth, with only vague 
cross-striolations finer than those of prozonite. 

Last tergite obtusely rounded behind, smooth. Anal valve 
with mesal valve strongly elevated, each set off by a broad 
furrow. Anal scale sharply set off by sulcus from its annulus, 
the sulcus evenly turved ; caudal margin only slightly convexly 
bowed out at middle. (See Pl. 13, Fig. 89.) Last annulus 
marked ventrally with strong cross-striations as shown in the 
figure. 


28 University of Michigan 
Number of segments, 55 to 57. 


Length, to near 80 mm.; width, 6 mm. 


Colombia: Fundacion River; Aug. 12, 1913. Four females, 
Holotype, M. C. Z., 5,090. 


Spirostreptus atoporus, sp. nov. 


Pl. 13, Figs. 90, 91 


While, in the absence of males, the narrower generic posi- 
tion of this species is doubtful, its characteristics are so pro- 
nounced that its recognition cannot give much difficulty, and 
it is accordingly here redescribed under Spirostreptus sens. lat. 
The fully developed color pattern shows a caudal ferruginous 
band about each metazonite, limited in front by a narrow, 
dark brown band passing into greyish-blue on anterior part 
of metazonite and on prozonite, the latter encircled also by a 
narrow brown stripe near edge of overlapping preceding meta- 
zonite. In some paratypes there is simply a ferruginous to 
fulvous caudal ring with a broader median dark brown to 
nearly black ring, and the covered part of prozonite again 
light, or the secondary darker stripes showing indistinctly. A 
middorsal dark line, and one also more or less evident on each 
side at level of pores. Collum bordered with dark, its median 
part areolated with light. Legs ferruginous. Four partly 
fused light spots between bases of antennae. 

Clypeal foveolae 2+2. Eyes rather small, the inner ends 
narrowly rounded or subangular, two and two-thirds, or a 
little more, their greatest diameter apart. Ocelli about 23, in 
four series; ¢. g., 7, 7, 5, 4; comparatively large. Outer face 
of cardo of mandible with caudal edge curving evenly into the 
distal one, the lower end angled only cephalo-distally. (See 
Pl. 13, Fig. go.) 


Occasional Papers of the Museum of Zoology 29. 


Lower end of collum but little inflexed; lower margin 
nearly straight; anterior angle well rounded, the posterior 
slightly obtuse. Six striae on each side above marginal one, 
these uniform and moderately fine and in part wavy. (See 
Pl. 13, Fig. go.) 

Covered region of prozonite only obscurely striolate,. 
excepting the stria most caudal in position, which is distinct. 
The suture is sharply impressed and smooth throughout. The 
pore is contiguous with the suture on its caudal side. Sur- 
face of segments in general seen under lens to be tubercular 
and in part obscurely rugose, the tubercles not close. Meta- 
zonites striate only beneath. 

Last tergite caudally obtusely angled. Anal valves with 
mesal borders only moderately elevated. Anal scale sharply 
set off from the annulus, conspicuously angled from behind, 
as shown in Pl. 13, Fig. 91; exposed portion of venter of last 
annulus smooth, not at all transversely striate. 

Number of segments, 60. 

Length, about 76 mm.; width, 4.5 mm. 


Colombia: San Lorenzo; July 2, 1913. One female in. 
jog at 4,500 feet elevation. Also one from a stump at same 
elevation, July 5, 1913. 

Cincinnati coffee plantation; July 2, 1913; A. S. Pearse. 
Two females under stone in cornfield. 

San Lorenzo; July 13, 1913; 4,500 feet. Two females. 
Also, July 14, 1913, one not fully mature male at 2,000 feet. 

One female without definite locality. 

Holotype, M. C. Z., 5,092. 

At once recognizable among the other spirostreptid species 
secured by the position of the repugnatorial pores and the 
weakly scabrous character of the surface of the segments. 


30 University of Michigan 


Orthoporus etholax, sp. nov. 
‘Pl. 14, Figs. 92-96 


This is a dark species characterized by having the caudal 
portion of the metazonites darker than the anterior portion, 
in specimens in full color the dark brown or dusky caudal 
band embracing a little more than a third of the length of 
metazonite and ordinarily bordered in front by an abruptly 
paler line or narrow stripe. Anterior region of metazonite 
and prozonite brown, the prozonite becoming paler cephalad, 
but with anterior border of covered portion ordinarily very 
dark. Last tergite dark excepting a narrow caudal border on 
each side, and sometimes median area somewhat lighter, this 
tergite and the anal valve typically of an olivaceous cast. Col- 
lum with a narrow light border all around limited within by 
a blackish band, the central area paler. Head dark across 
vertex, paler below. Legs typically dark brown to light brown, 
the legs agreeing in color in general with the body. 

Vertigial sulcus of head distinct. Clypeal foveolate 2+2. 
Eyes transversely elongate, with inner end acute and upper 
margin convex; separated by less than their diameter. Ocelli 
arranged in six series, about 56 or 58 in number; e. g., 13, 12, 
11,9, 7, 4, and 12, 11,9, 8,5, 2. Head back of antennal socket 
and below eye with a smooth margining ridge, this ridge con- 
tinuing along the notch; area in front of the ridge roughened. 

Collum in female with inferoanterior angle obtuse, the 
inferocaudal one rounded. Typically with three deep sulci 
above the margining one and a characteristic furrow curving 
dorsocaudad from near middle of uppermost of these sulci, 
this furrow not attaining the caudal margin. Or there are 
four sulci if the weaker one at edge is regarded as margining 
sulcus. (See Pl. 14, Fig. 93.) Collum of male with ante- 


Occasional Papers of the Museum of Zoology #1 


ventral corner produced ventrocephalad, rounded distally. 
(Pl. 14, Fig. 92.) | 

Segmental sutures sharply impressed throughout, smooth, 
curved opposite the pore, which is less than half the distance 
from the suture to the edge of the caudal dark band of meta- 
zonite. Metazonite striate only beneath, but with same obscure 
broad and very low rugae above the striae proper. 

Last tergite with caudal portion set off by a very shallow, 
more or less obscure, transverse depression or furrow; caudal 
angle acute. Mesal borders of anal valves compressed and 
elevated, the elevated region of only moderate height. Anal 
scale sharply set off; each side concave, the caudal angle rec- 
tangular or a little obtuse. (PI. 14, Fig. 94.) 

Gonopods of male as shown in Pl. 14, Figs. 95, 96. 

Number of segments, 54 to 55. 


Length, about go mm.; width, 6.5 mm. 

British Guiana: First Mourie. In Calladium root masses 
in tree clump. Aug. 1 and 5, 1914; F. M. Gaige. Two females. 
Labba Creek Sand Hills; July 27, 1914. One female. 

Forested Sand Hills; Aug. 18, 1914; F. M. Gaige. One 
male in Calladium root mass. 
Holotype, M. C. Z., 5,098. 


Orthoporus walkeri, sp. nov. 


Pls 74s Pic. 07> Pl 15; Big, 68 


Somewhat resembling the preceding species, O. etholax, 
but readily distinguished by difference in color pattern as well 
as in details of structure. Unlike etholax, this species has a 
caudal band of each typical segment lightest, subferruginous 
in color with the anterior portion of metazonite, embracing 


more than half of it, deep brown of olive cast. Last tergite, 


32 University of Michigan 


excepting a narrow caudal border on each side and anal valves 
nearly black or black of olive tinge. Legs brown, lighter than 
in ectholax, 

Most easily separated from etholax by readily noted dif- 
ferences in the collum. The inferoanterior corner in the 
female is not obtuse, but is moderately extended forward as 
shown in Pl. 15, Fig. 98. The collum lacks the oblique fur- 
row above the striae proper, and the next to the uppermost 
stria is short, crossing only the caudal border as shown in the 
figure. Five striae are present above the margining sulcus. 

Segmental suture deeply impressed, widely curving oppo- 
site the pore, which is a little in front of a point halfway 
between suture and caudal edge of dark portion of metazonite. 

In the type there are two transverse furrows in front of 
the tip of the last tergite. Anal valves with mesal borders 
elevated as usual. Anal scale longer in proportion to width 
than in etholax. (PI. 14, Fig. 97.) 

Number of segments, 61. 

Length, about 95 mm.; width, 6.5 mm. 

British Guiana: First Mourie; July 30 and Aug. I, 1914; 
F, M. Gaige. One female on each date. Holotype, M. C. Z., 
5,101; 

Orthoporus gaigei, sp. nov. 


P]. 15, Figs. 99-103; Pl. 16, Fig. 104 


Segments typically mostly blackish brown or black, except- 
ing a narrow ferruginous ring about caudal border of each 
and the covered por‘ion of prozonite, which is also pale. Ante- 
rior border of anal valves typically palest. Head and collum 
often in large part chestnut or ferruginous. Antennae and 
legs ferruginous. 


Vertigial sulcus of head sharply impressed, ending in a 


Occasional Papers of the Museum of Zoology 33 


depression or pit at level of inner angles of eyes, where it 
meets a vaguer transverse line from each side. Clypeal foveo- 
lae 2+2. Eyes angled within, separated by about two-thirds 
their length. Antennae, when laid back on dorsum parallel 
with median dorsal line, not fully attaining the caudal edge 
of the dorsum; when laid back along the side a little surpass- 
ing the caudal margin of collum. 

In the female the lower edge of the collum on each side 
is nearly straight with the caudal angle widely rounded, the 
cephalic angle subrectangular, narrowly rounded. (See PI. 
15, Fig. 99.) In the male the anterior angle of collum is pro- 
duced, as, e. g., in the male of Nanostreptus incertelineatus 
Silv., but more forward, the end of process rounded. (See 
Pl. 15, Fig. 100.) On each side below there are, in addition 
to the margining sulcus, three complete sulci, with sometimes 
(in females, at least) one or more less complete intermediate 
ones. 

Covered portion of prozonites with encircling striolae that 
branch and anastamose in the ordinary manner. Exposed por- 
tion of prozonite and the metazonite subdensely finely punc- 
tulate, otherwise smooth and shining above and laterally, the 
metazonites longitudinally striate beneath. Metazonites not 
elevated. Suture sharply impressed throughout, only slightly 
and widely curving opposite pore, which is widely removed 
from it, lying approximately on the line between first and 
second thirds of metazonite. 

Last tergite with a subtriangular caudal region set off by 
a shallow transverse furrow, the angle rounded; with no 
median keel. Anal valves with mesal borders strongly com- 
pressed and sharply elevated. Anal scale separated from the 


annulus by a deep sulcus ; broadly triangular. (PI. 16, Fig. 104.) 


34 University of Michigan 


Gonopods of male as shown in Pl. 15, Figs. 101-103. 

Number of segments, male type, 58; female, 57 to 62. 

Length, female, to near 150 mm.; width, 8 mm. Length 
of male type, about 110 mm.; width, 7 mm. 

Colombia: San Lorenzo. Under leaves, 2,000 feet to 
3,000 feet. July 16, 1913; F. M. Gaige. One male and four 
females. Also along trail, between 2,500 and 3,000 feet. July 
7, 1913; M. A. Carriker. Two males. At 2,000 teet> julg 
24, 1913; one male. 

Colombia: San Lorenzo. At edge of clearing; 4,500 feet; 
July 9, 1913. A female and a variant male. At 4,500 feet; 
in leaf-cutter ant’s nest; one female. July 16, 1913. 

Holotype, M. C. (Z., 5103. 


Orthoporus foliatus, sp. nov. 


Pl. 16, Fig. 105 


General color deep olive brown. In the more anterior seg- 
ments there is a very distinct light fulvous to fulvoferruginous 
band about caudal portion of metazonite, the extreme caudal 
border of which, however, is darkened. ‘The light band becomes 
somewhat less distinct going caudad; in middle region on 
dorsum it is obviously less than one-third the total length of 
metazonite. Legs reddish brown. 

Vertigial sulcus distinct. Eyes in type with ocelli in 
eight series, 83 in number; ¢. g.,°15, 14, 14, 11, 11, Tipe 
Area behind antenna and below eye characterized by having 
a rather broad inferior border which is smooth, the region 
above it roughened by an oblique ruga or ridge from each 
side of which others extend like the veins of a leaf from the 
midrib. (See Pl. 16, Fig. 105.) 


The lower anterior angle of the collum is carried some- 


Occasional Papers of the Museum of Zoology 35 


what forward as in O. walkert. There are only four striae 
or sulci above the lowest margining one on each side. Of 
these, the second one is characteristic, being parallel to margin 
and ending abruptly in front, not curving and continuing 
upward as do the others. (See Pl. 16, Fig. 105.) 

Segmental suture deep, as usual, a little excurved oppo- 
site the pore which lies in about the middle of the darker 
region of metazonite. Metazonites of middle segments striate 
only well beneath; but up the sides and across dorsum rough- 
ened by numerous somewhat irregular and very weak longi- 
tudinal rugae or folds which may, with proper lighting, be 
seen with the unaided eye. 

Last tergite characterized by having the caudal portion 
sharply set off by a transverse furrow and depressed below 
level of anterior region; also more strongly roughened, finely 
reticulo-rugose, just caudad of the furrow. Tergite rugose 
on the sides. Anal valve with mesal borders elevated as usual. 
Anal scale of usual general form, rather broad, with a broad 


median longitudinal elevation on keel. 
Number of segments, 58. 
Length, about 105 mm.; width, 7.5 mm. 


British Guiana: Sand Hill Forest. In Calladium root 
mass. Aug. 22, 1914; F. M. Gaige. One female. Holotype, 
mae. Z., 5,100. 


Orthoporus sp. 


In the collection are several immature females and males 
of an uncertain species. 

British Guiana: Forested Sand Hills. Aug. 17 and 18, 
1914; F. M. Gaige. 


36 University of Michigan 


SPIROROLOIDEA 


RHINOCRICIDAE 


Rhinocricus brevipes Carl 


Colombia: San Lorenzo. Under leaves at elevation of 
2,500 feet, July 15, 1913; and at elevation of 2,000 feet, July 
16, 1916; F. M. Gaige. , 

Four females agreeing well in all structural details with 
the original description, excepting that the segments number 


A4, 45, and 47, instead of 42. 


Rhinocricus arboreus Saussure 


Unted States Virgin Islands: St. Croix; Sept. 14-18, 1914. 


‘ 


Mr. Gaige, the collector, notes that these millipedes “were 
abundant over the island, usually found in the trees, where 
they frequently gathered in clusters of three or four to twenty 
or more.” 
Rhinocricus monilicornis (Porat) 

United States Virgin Islands: St. Croix; July 16, 1914; 
F. M. Gaige. Four specimens taken with Trigoniulus lum- 
bricinus Gerstaecker in interior of island. 

British Guiana: Labba Creek Sand Hills; July 27, 1914; 
F. M. Gaige. Two males and six females collected in rotten 
wood. 

East Trail along Demerara River, close to high-water 


mark; Aug. 8, 1914; F. M. Gaige. One male and one female. 


Rhinocricus hylophilus, sp. nov. 
Female——The body in general is deep olive to nearly black, 
excepting the caudal borders of metazonites, which are dull 
ferruginous forward more than half way to the suture, and 
the covered portion of prozonites, which is paler, as usual. 


Collum bordered with ferruginous. Head on the lower bor- 


Occasional Papers of the Museum of Zoology 37 


der of face ferruginous. Antennae and legs also ferruginous, 
excepting the two proximal joints, which are darker, more 
brownish or olivaceous. 

Sulcus of head interrupted in the interocular region. Cly- 
peal foveolae 2+2. Antennae clavate; sensory cones numer- 
ous. Facets of eyes flat, often obscure; eye subtriangular, with 
apex ectad; ocelli about 28 in five transverse series; e¢. g., 7, 7, 
6, 5, 3; the ectal ocelli of upper series much the largest, the 
others decreasing mesad and centrad. Eyes nearly three 
times their greatest diameter apart. 

Collum widely rounded on each side below, the second ter- 
gite extending considerably below its ends; not margined, stri- 
ate or distinctly punctate, being smooth and shining. 

In a typical segment from the middle region of the body 
the suture is fine but complete, angled at level of pore which 
it touches, the suture lying in a furrow which is vague dor- 
sally but becomes distinct below level of pore. No secondary 
furrow. A little in front of the suture a shallow furrow, 
which is often irregularly developed and may be evident only 
across dorsum above level of pores, its lower end on each side 
often curving cephaloventrad. Covered portion of prozonite 
with the usual transverse sulcus across dorsum just caudad 
of this striolate region. Metazonites with deep longitudina 
striae only below, the non-striate region between pore and the 
striae twice as wide as the striate region. Under the lens this 
region between striae and pore shows weaker short impressed 
lines which are in part branched and scattered punctae, this 
same sculpture continuing in a band caudad of the suture up 
some distance above pore on each side, the metazonite other- 
wise wholly smooth. 

Scobina distinct to segment 22 or 23, at which they cease 


abruptly. 


38 University of Michigan 


Last tergite with cauda acute, straight, extending a short 
distance beyond the valves, a transverse sulcus at base of 
cauda. Valves with mesal borders strongly compressed and 
elevated. Anal scale smooth. 

Number of segments, 44 to 46. 

Length, near 155 mm.; width, 18 mm. 

Colombia: San Lorenzo. On trail in heavy forest, between 
2,000 and 4,500 feet elevation. July 1, 1913. Three females. 

Poletyne, MM. CZ.; 5, Tro, 


Rhinocricus pycnus, sp. nov. 


Female.—Dark olive, the segments bordered with ferru- 
ginous behind, the ferruginous band extending nearly to the 
suture. Last tergite narrowly bordered behind with ferrugi-’ 
nous and the elevated mesal margin of valves the same color. 
Collum bordered with ferruginous. Clypeus ferruginous, this 
color also extending up in the middle between the antennae, 
but here more obscure. Eyes also ferruginous. Legs and 
antennae ferruginous. 

Sulcus of head interrupted at level of eyes. Clypeal fove- 
olae 2+2. Antennae flattened, of uniform width over most 
of length; sensory cones numerous. Eyes consisting of about 
28 ocelli in five series; e. g., 7, 7, 6, 5, 5; facets distinctly con- 
vex, decreasing in size mesad, as usual. 

Collum only slightly exceeded on each side by the second 
tergite; ends rounded; surface without margining or other 
sulci, under the lines showing a sculpturing of fine coriarious 
markings. The suture on typical segments of body complete 
and distinct, slightly angled at level of pore which it touches. 
Longitudinal striae of metazonite few, those present present 
only below. Elsewhere metazonite and exposed portion of 


prozonite seen under lens to be covered with numerous 


Occasional Papers of the Museum of Zoology 39 


impressed fine lines which branch and in part anastomose like 
those on the collum. Covered portion of prozonite with the 
usual wavy transverse striolations. 

Scobina ending on the twenty-third segment. 

Last tergite without transverse sulci; with an acute cylin- 
drical cauda produced well beyond the valves and distally a 
little curved upward. Anal valves with mesal margins com- 
pressed and elevated as usual. Anal scale smooth, caudal 
angle obtuse, sides convex caudally and straight anteriorly. 

Number of segments, 43 to 44. 

Length, 54 mm.; width, 8.5 mm. 

Colombia: San Lorenzo. Among fallen leaves at 2,000 
feet, July 14, 1913, and under leaves at 4,200 feet, July 15, 
1913. One female at each locality. Holotype, M. C. Z., 5,112. 

The body of this form is usually thick in proportion to its 
length. 


Rhinocricus amblus, sp. nov. 
Pl. 16, Figs. 106-108 


Body dark, the exposed portion of the prozonites dark 
bluish to brownish, the metazonites mostly abruptly darker, 
deep brown, excepting a narrow, pale caudal border. Covered 
portion of prozonites pale, with a network of darker lines. 
Segments often paler below adjacent to legs. Head ferrugi- 
nous excepting for a dark subquadrate mark between the eyes, 
the vertex covered with a network of dark lines and a similar 
network at the sides of and below the dark mark. Collum 
sometimes ferruginous, with network of darker lines. Legs 
and antennae ferruginous. Last tergite and anal valves dark, 
excepting a narrow, pale border on each side of the tergite, 
the cauda wholly dark. When dry the body may appear essen- 
tially black excepting for the pale caudal border of each 
segment. 


40 Unizersily of Michigan 


Median sulcus of head interrupted at level of eyes. Cly- 
peal foveolae 2+2. Antennae very short, with only four sen- 
sory cones. Eyes small, subcircular. 

Collum rounded at each end, with a short margining sulcus 
across the end on each side. Surface densely but very finely 
punctate and punctolineate. Second tergite extending a little 
below end of collum on each side. The typical segments of 
body have the suture sharply impressed throughout, the seg- 
ments a little constricted at its level; suture bending angularly 
a little away from pore at level of latter. Pores minute. Cov- 
ered portion of prozonite smooth. Exposed portion of prozo- 
nite anteriorly, with two fine transverse striae across dorsum. 
The prozonite and metazonite in general with numerous 
minute impressed points and short lines running out from 
these points as frequent in Trigoniulus, these more marked 
on the prozonite than on the metazonite. Segments longitudi- 
nally sharply striate beneath. 

Last tergite smooth, with a subcylindrical cauda which is 
rounded at the end and extends beyond the anal valves. (PI. 
16, Figs 106, 107.) Anal valves not margined, the mesal bor- 
ders rounded, slightly compressed, but not set off. Caudal 
margin of anal scale convex. 

For gonopods of male, see PI. 16, Fig. 108. 

Number of segments: female, 45 to 46; male, 43. 

Length, 42 mm.; width, 4.7 mm. 

Colombia: San Lorenzo. Under leaves and logs at 2,500 
feet; July 15, 1913. Four females. At 4,500 feet, in stumps; 
July 5, 1913. Three females. 

Under log, 4,500 feet, July 2, 1913. One female. 


Under log, 4,500 feet, July 3, 1913. One male. F. M. 
Gaige. 


Holotype, M. C. Z., 5,114. 


Occasional Papers of the Museum of Zoology 41 


SPIROBOLELLIDAF, 
Microspirobolus tridens, sp. nov. 


Pe. 76. Rie too: Pl.-ez Bigs 110 


Dorsum fulvous to buff, with a middorsal stripe formed by 
a series of black triangular marks with bases cephalad, the 
lateral angles extending laterad in black lines along anterior 
borders of the somites to unite with the black band along each 
side. Venter and lower part of sides again light colored 
excepting borders of segments. Last tergite black, excepting 
a light spot anteriorly on each side. Collum also blackish, 
excepting a pale spot on each side behind. Legs ferruginous. 

Eyes about equal in length and breadth. Ocelli about 18 
in only four series; @. g., 5, 5, 5, 3. Eyes separated by a space 
three times as long as their diameter. Antennae very small. 

Collum narrowed moderately down each side, the lateral 
ends rounded and bent a little mesad below; margin below 
and up the front as usual. 

Each ordinary segment with a moderate encircling furrow 
or constriction; this crossed by numerous ridges or ribs both 
on dorsum and down the sides, the depressions between the 
ridges on the sides continuing upon the prozonite as deep 
striae which curve upward. Metazonite striate below. 

Last tergite rounded caudally, projecting caudad freely 
above valves for a short distance. Anal valves weakly mar- 
gined along mesal edges, the borders not protruding. 

Characterized especially by the form of the gonopods of 
the male, which are represented in Pl. 16, Fig. 109, and PI. 
e7, Higs-110. 

Number of segments, 34 to 37. 

Length, about 18 mm.; width, to 2 mm. 

Colombia: San Lorenzo; 4,500 feet; July 3, 1913. Four 


42 Umiersity of Michigan 


males and three females taken under bark of stump. Holo- 
type, BEAC. Zz S118. 


TRIGONIULIDAE 


Trigoniulus lumbricinus (Gerstaecker ) 


United States Virgin Islands: St. Croix; July 16, 1914; 
F. M. Gaige. “Collected in rotten logs in deep ravine in inte- 
rior of island. This form was a very abundant one, and was 
present under almost every stone that was turned over, as 
well as in rotten wood, beneath loose bark, etc. It was noted 
to be equally abundant on the high and very dry hillside and 
in the damp ravine.” In the vial with the Trigoniulus were 
four specimens of Rhinocricus monilicornis (Porat), to which 


this note in part, therefore, applies. 


POLYDES MOIDEA 
EURYURIDAE 
Pycnotropis colombiensis, sp. nov. 
Bis sess iT us 

When in full color the dorsum is blackish brown to black, 
with the keels fulvous. Last tergite paler, brown or reddish 
brown, across caudal border. Legs and antennae brown. 

Vertigial sulcus sharply impressed, lying in a deep furrow 
which anteriorly branches, sending a_ shallow furrow 
obliquely out to each antennal socket. 

Collum narrower than the succeeding tergite. Lateral 
lobes or keels wholly smooth, conspicuously narrowed and the 
ends strongly convex. Surface between the keels divided into 
polygonal areas like those of other tergites, these areas form- 
ing four transverse, in which the areas of the first and last 
are more or less incomplete. (Pl. 17, Fig. 111.) 


The polygonal areas of the tergites in general more pro- 


Occasional Papers of the Museum of Zoology 43 


nounced toward the keels, with their surface more convex 
and the tubercles correspondingly more distinct. 

In the last tergite the cauda has the caudal margin convex 
and crenate, while the lateral margins are essentially straight 
and smooth, excepting a single setigerous notch. 

Sternite of fourth segment anteriorly, between fourth legs, 
with two straight conical processes, and with two smaller con- 
ical processes between fifth legs. The sternite of the fifth 
segment also bears two pairs of similar processes, of which 
those of the posterior pair are much more widely separated 
than those of the anterior pair. 

In the gonopods of the male there is near the middle of 
the principal distal branch an acute tooth below, and one or 
sometimes two smaller teeth proximad of the acute distal end. 
(See, further, Pl. 17, Figs. 112 and 113.) 

Length of male, about 42 mm.; width, 6.8 mm. 

Colombia: San Lorenzo. Four specimens taken under a 
log at 4,500 feet, July 2, 1913. Ten specimens, partly imma- 
ture, on logs at 3,000-feet, July 16, 1913. One male and two 
females in stumps at 4,500 feet, July 4, 1913. F. M. Gaige. 

Cincinnati Coffee Plantation. One male and three females, 
one immature, taken in a rotten log, July 2, 1913. A. S. 
Pearse. 

Holotype, M. C. Z., 5,120. 


Pycnotropis cylindroides, sp. nov. 
Pl. 17, Figs. 114-116 


Dorsum a dull or grey olive brown throughout, the keels 
not lighter in color, or somewhat more reddish than dorsum; 
the olive cast sometimes not apparent. (Male type.) Anten- 
nae light brown. Legs yellow to light brown of slight olive 
tinge. 


44 University of Michigan 


Contrasting strongly with the previous species not only in 
color but also in form from the great narrowness of the carinae, 
this giving the body a nearly cylindrical appearance. The 
carinae in general are so narrow that the poriferous swelling 
occupies the entire width in posterior part, with the part in 
front still narrower, merging into the general surface of the 
segment. Anterior keels wider than the posterior ones. Pos- 
terior angles slightly produced from fifth segment caudad. 
Dorsal surface of segments divided into polygonal areas as in 
P. colombiensis, but tubercles absent or obsolete and irregu- 
larly developed. Some metazonites more or less roughened 
toward the keels. See further, Pl. 17, Figs. 114 and 115. 

Caudal margin of last tergite strongly convex, the sides 
converging caudad. 

Gonopods of male as shown in Pl. 17, Fig. 116. 

Length, about 45 mm.; width, 6.5 mm. 

Colombia: San Lorenzo. A female in its “mud nest” with 
eggs was taken under a log at 5,300 feet elevation, and a male 
under a log at 5,000 feet, July 23, 1913. Holotype, M. C. Z., 
Bi25. 

PLATYRRHACIDAE 
Rhyphodesmus amphelictus, sp. nov. 
Pl. 18: Sbigs. Li7e iro 


General color dark brown, the keels scarcely paler, but the 
cauda over all but anterior portion abruptly lighter. 

Surface of head densely granular, with granules of differ- 
ent sizes. A deep furrow across vertex, widening anteriorly 
to occupy entire space between the rims of antennal sockets. 

Collum much narrower than the second tergite (width 
about as 7.25 to 10); lateral end in general outline subrectan- 


gular, but a little rounded at apex. Surface densely granular, 


Occasional Papers of the Museum of Zoology 45 


a row of larger tubercles along anterior and posterior margins 
and tubercles of similar size in intervening space not serially 
arranged; on each side an area of granules or tubercles ele- 
vated above margin proper. 

Each ordinary metazonite with dorsal surface densely 
granular and with three transverse series of larger tubercles, 
the three series equally developed. Along the lateral margin 
of the keel a series of much larger rounded tubercles giving 
the appearance of a fine or less even serration. Pore removed 
from margin by a distance of from one and a half to two 
times the diameter of its rim. 

Cauda subquadrate, broader than long, a little narrower 
at base than in middle of its length, the sides and the caudal 
margin being weakly convex. 

Gonopods of male with two dista! branches, these smooth 
and unbranched, the lower (posterior) one much shorter, 
curling dorsad distally, the larger branch curling into a circle 
and in situ resting against the sternite of the sixth segment. 
mee further, Pl. 18, Figs. 117,118. 

Length, male, 70 mm.: width, 12 mm. 

British Guiana: Labba Creek Sand Hills; July 27, 1914; 
F. M. Gaige. One male and one female. 

Two males and two females taken at Cacao Plantation 
about camp in rotten wood, July 16 and 17, 1914; F. M. Gaige. 

One adult and several immature specimens from bank of 
Demerara River collected in a very damp and much decayed 
stump; July 17, 1914; F. M. Gaige. 

Eielotype, M:C.°Z., 5,127. 

I adopt for the genus of this species the name Rhyphodes- 
mus, although the species on which Cook based his brief diag- 
nosis of the genus has not as yet been described and cannot 
be determined from the published data. 


46 University of Alichigan 


Aphelidesmus guianensis, sp. nov. 


Pl. 18, Figs. 119-122 


Dorsum with metazonites deep colored or blackish, the 
prozonites in some degree lighter and the keels entirely pale 
fulvous or, in types as preserved, nearly white. Last tergite 
dark throughout. Legs and antennae light brown. 

Collum strongly narrowed laterad, the lateral lobe or keel 
near its outer end only one-third as long as the middorsal line 
of the plate, while at its base, 7. e., near inner end of light 
area, about two-thirds as long as that line. Anterolateral cor- 
ner widely convex, the caudolateral more narrowly rounded. 
(Pl. 18, Fig. 119.) 

In the second tergite the lateral margin of keels straight, 
with the two angles in general outline rectangular, but the 
outer edge of the anterior one produced ectad. In the third 
tergite the caudolateral angle of keel moderately acute, the 
outer edge of keel oblique and a little convex, the anterolat- 
eral corner with ectal process smaller. In the fourth tergite 
the outer edge of keel still more convex, with anterior corner 
more rounded and the process or tooth still slighter and 
rounded, scarcely evident, as shown in PI. 18, Fig. 120. 

Cauda with sides convex; caudal margin obtusely notched 
at middle. Anal scale very broadly shield-shaped, the sides 
convex excepting at anterior end, where concave; posterior 
angle very obtuse. (PI. 18, Fig. 121.) 

In the male gonopods the tarsus characterized by bearing 
a short, plate-like process on its caudal side. See further, 
Pl. 18, Fig. 121. 

Length of male type, 46 mm.; width, 7 mm. 

British Guiana: Forested Sand Hills; Aug. 17, 1914; F. 


Occasional Papers of the Museum of Zoology 47 


M. Gaige. One male and two females collected in rotten wood 
and in the earth. 

Sand Hill Forest. One male in rotten wood, Aug. 19, 
1914, and two females m Calladium root masses, Aug. 24, 
1914; F. M. Gaige. 

First Mourie. One female “collected in ground litter in 
tree clump,” Aug. 6, 1914; F. M. Gaige. 


Polylepiscus sp. 
One young and badly broken specimen of this genus was 
secured. Its poor condition renders it inadvisable to attempt 
to place or describe it specifically. 


Colombia: San Lorenzo; Cincinnati Coffee Plantation. 
In a rotten log; July 2, 1913; A. S. Pearse. 


LEPTODES MIDAE 
Trachelodesmus angulatus, sp. nov. 


Pl. 18, Figs. 123-125 


Metatergites greyish blue, the prozonites paler. Sides just 
under the keels also greyish blue, the lower part of sides and 
the venter fulvous. Head with ground yellow ; a network of 
dark lines over vertex and darker areas below this and in 
region below level of antennae. Collum also with a network 
of dark lines. Legs and antennae fulvous. 

Body constricted anteriorly, as usual. 

This species stands apart from others thus far described 
in the details of the metazonites, as notably in the acute, cau- 
dally produced caudal angles of all keels from the fifth caudad 
and the second to fourth also bearing at the caudal angle a 
caudally directed but less conspicuous tubercle, this uniform 
with the tubercles of the caudal series. Dorsal surface of 


metazonites densely granular, with each typical one divided 


48 University of Michigan 


into three transverse rows of large polygonal areas, of which 
the most caudad are incomplete, only the anterior halves being 
present; six areas in each series. From caudal edge of meta- 
zonite project six larger tubercles directly caudad, one tubercle 
from each area. Each area of the two anterior rows also 
with a less conspicuous central tubercle among the smaller 
granules. Tubercles of margin of keels larger, more or less 
directed caudad of ectad, serriform, the most caudal largest, 
particularly large on the poriferous segments on which it is 
confluent, with one or two preceding tubercles to form the 
large poriferous prominence on which the pore opens below 
the upper edge, a nodule, the tip of a fused tubercle, lying 
just over it. (See Pl. 18, Fig. 124.) Sides obscurely granu- 
lar. Second, third and fourth tergites with three marginal 
tubercles or teeth, of which the one at the anterior corner is 
smallest ; the caudal series of tubercles as in the more caudal 
segments, but only one series in front of this. Legs not 
granular. 

Collum acutely angular on each side, a tubercle at apex, 
margin convex each side of this angle. Six tubercles along 
caudal border, and two rows of rather obscure tubercles far- 
ther forward, the surface otherwise densely granular. (Pl. 
18, Fig. 123.) 

Sternal spines present larger than those of corresponding 
segments in other species, but the most caudal ones not larger 
than the others, all being curved and subequal excepting those 
of eighth and ninth segments, the most anterior ones present, 
which are reduced. (See Pl. 18, Fig.’ 125.) 

Length, about 16 mm.; width, 2.5 mm. 

Colombia: San Lorenzo, Two females of 19 segments 
and two younger specimens taken in forest at 4,000 feet; July 
14, 1913; F. M. Gaige. Holotype; M. C. Z., 5,136. . 


Occasional Papers of the Museum of Zoology 49 


Trachelodesmus ancylophor, sp. nov. 


Pl. 19, Figs. 126, 127 


General color light horn-brown, a considerable portion of 
metazonite and prozonite above darker, dusky, in the type the 
collum and vertex of head with a network oi dark lines. Legs 


and antennae fulvous. 


Body rather slender, widest near middle and narrowing 
very gradually forwards, the anterior constriction including 


particularly the third and fourth segments. 
The head not granular or tuberculate, smooth throughout. 


Collum a little narrower than head, inclusive of cardines 
of mandibles. Margin widely and evenly convex over ante- 
rior and lateral regions back to caudolateral angles, which are 
subrectangular ; caudal margin convex on each side just within 
the angle, the median region nearly straight. The dorsal sur- 
face presents along the caudal border about ten larger, more 
elevated, rounded tubercles, between which are wedged in 
smaller tubercles forming in general two rows, one in line 
with anterior and one with posterior borders of the larger 
tubercles ; the remaining portion closely covered with smaller 
and lower flattened tubercles back of anterior border, such 
as occur in Trachelodesmus constrictus Peters. A row of 
about six widely separated setae just behind the depressed 
anterior border. (Pl. 19, Fig. 126.) 

The second tergite with two irregular series of mostly 
large tubercles across median region; the principal tubercles 
on each side are much larger than those of median region, 
there being behind them a caudal series of much smaller 
tubercles ; a large, blunt tubercle at anterolateral corner is car- 


ried forward against the end of the collum. Third tergite 


50 University of Michigan 


also with two rows of rounded tubercles, the fourth with an 
irregular series of smaller tubercles between the other two, 
two large, rounded tubercles on lateral margin of keel. A 
typical metazonite of middle region of body has the dorsal 
surface divided into large polygonal areas, which are more 
distinct toward the sides, these forming two transverse rows 
of mostly eight areas each; these larger areas bear densely 
arranged smaller tubercles. Lateral margins of poriferous 
metazonites convex, each with a very large, rounded tubercle 
through the ectal face of which the pore opens. Suture 
between metazonite and prozonite longitudinally striate as in 
constrictus. Prozonites finely granular above. (PI. 19, Figs. 
126, 127.) 

Cauda triangularly narrowed caudad, narrowly truncate 
at tip; caudal surface bearing six setae; two setae on each 
side and a pair above, in front of a more widely separated 
pair on basal portion of tergite. 

Sterna very wide, as usual, but narrower than the tergites 
exclusive of keels; not granular or tuberculate. Sterna from 
thirteenth to eighteenth bearing small spines in the usual posi- 
tion adjacent to the posterior legs, the nineteenth segment with 
a pair of very much larger curved spines. 

Sides of segments smooth, or nearly so. 

Legs not granular, or only very obscurely so, but other- 
wise typical in form. 


Length, about 15 mm.; width, 2 mm. 

Colombia: San Lorenzo. One female under a log at about 
6,200 feet, July 19, 1913. 

One broken female at 6,200 feet, July 19, 1913, and two 


females from a locality of which the number cannot longer 
be deciphered. 


Occasional Papers of the Museum of Zoology 51 


One female among bromeliads on ground at 7,600 feet, 
July 19, 1913. 

Holotype, M. C. Z., 5,138. 

Resembles T. constrictus Peters, but differs—e. g., in lack- 
ing the anterior row of larger tubercles on the collum, in 
having two rows of tubercles, instead of but one, on the sec- 
ond tergite, in having sternal spines only from thirteenth seg- 
ment caudad, and conspicuously in having the sterna and legs 


free from true granulations. 


Dromodesmus, gen. nov. 

Body composed of head and 20 segments. 

Sternum of nineteenth segment narrowed caudad, the pos- 
terior pair of legs being rather close together and projecting 
directly caudad, or nearly so. No spines on this sternum, but - 
preceding ones of posterior and middle segments with a pair 
of spines at bases of posterior pair of legs. 

Legs exceptionally long. 

Keels of anterior and median regions well developed, car- 
ried high, margins smooth, and posterior angles of all from 
the second caudad conspicuously produced ; the posterior keels 
reduced anteriorly, leaving only the caudolateral processes 
prominent, the posterior end of body narrowing conspicuously 
caudad. Dorsal surface of metazonites densely granular and 
with longer tubercles in transverse series. Repugnatorial 
pores on segments 5, 7, 9, 10, 12, 13, 15-19. Cauda triangular, 
caudal apex narrowly truncate. 

Gonopods of male with telopodite deeply bifid, both 
branches segmented, the outer (anterior) toward base, the 
inner (posterior) toward middle; outer branch in genotype 
trifid, with two processes acute and one plate-like; the inner 
branch distally bifid. 

Genotype, Dromodesmus longtpes, sp. nov. 


52 University of Michigan 


Dromodesmus longipes, sp. nov. 


Pl. 19, Figs. 128-133; Pl. 20, Figs. 134-135 


The general color of most of the types as at present is 
brownish grey, with legs fulvous and antennae brown. But 
the only specimen in full color, the male type, is deeply olivea- 
ceous or nearly black above, excepting the processes or the keels, 
which are yellow. Lighter, brownish yellow beneath. Anten- 
nae brown. Legs lighter brown, fulvous beneath proximally. 

Antennae long (see Pl. 19, Fig. 131, for proportions of 
joints). Surface of head minutely granular, with fewer 
longer setigerous granules. Vertigial sulcus distinct to level 
of antennae. 

Collum with anterior and lateral margins together forming 
a nearly evenly convex curve. Caudal angles a little less than 
rectangular, extended slightly caudad. (PI. 19, Fig. 128.) 

Keels of anterior segments bent up and carried a little 
above level of dorsum, in going caudad becoming more nearly 
horizontal. All margins of keels wholly smooth. Dorsum 
of a typical metazonite with a distinct transverse furrow in 
front, of which there is a median longitudinal furrow, often 
irregularly developed, giving a number of quadrangular areas; 
a row of smaller areas along caudal border. Two transverse 
rows of tubercles caudad of the transverse suture and one in 
front, the latter of six widely separated tubercles. (For form, 
see Pl. 19, Fig. 129.) 

Cauda triangular, with tip narrowly truncate; a large 
setigerous tubercle on each side, with a smaller one more 
dorsal in position between it and the tip. Mesal margins of 
valves raised. Anal scale angular behind, with a setigerous 
tubercle each side of the angle. (PI. 19, Fig. 130.) . 

Form of posterior legs of male as shown in PI. 20, Fig. 134- 


Occasional Papers of the Museum of Zoology 53 


Gonopods of male as shown in PI. 20, Fig. 135, and PI. 19, 
Fig. 132. 

Length, female, 36 mm.; width, 5 mm. 

Colombia: Fundacion. About 18 specimens, mostly badly 
preserved and faded, from under a log along Fundacion 
River, Aug. 12, 1913. Holotype, M. C. Z., 5,142. 


Colombodesmus, gen. nov. 


Like Trachelodesmus and related genera in having sterna 
broad and in large part bearing paired spines, but a spine at 
base of each leg on segments where present instead of only 
at base of legs of each posterior pair, and the last pair of 
spines not more strongly developed than the others. Keels 
well developed in anterior region, marginally serrate, becom- 
ing more weakly developed, less projecting in middle and pos- 
terior region. The pores borne on distinct lateral processes 
on caudal portion of margin; present on segments 5, 7, 9, I0, 
12, 13, 15-19. Body not obviously constricted in neck-line form 
anteriorly. Metatergites finely tubercular or granular. Cauda 
triangular, tip narrowly truncate. Legs, in the genotype at 
least, granular, as in most species of Trachelodesmus; the 
claws well developed. 


Genotype, C. catharus, sp. nov. 


Colombodesmus catharus, sp. nov. 
Pl. 20, Figs. 136-140; Pl. 21, Fig. 141 


The type at present has the prozonite above and on the 
sides smoky brown, the body otherwise at present light grey 
throughout excepting a pair of light brown spots on the front 
of metatergites, the two spots on collum large and subconflu- 
ent. Legs and antennae light yellowish grey, darker distally, 
especially the tip of antennae. 


on 
ars 


University of Michigan 


Antennae inserted not far apart, the distance between their 
sockets but little greater than the diameter of the latter. Head 
depressed or widely furrowed from base of each antenna cau- 
dolaterad, the proximal portion of antenna lying in this fur- 
row. Vertigial sulcus distinct to level of antennae. Setae 
over entire surface of head numerous and nearly uniformly 
distributed. 

Collum but little narrower than head inclusive of cardines. 
Anterior margin convex, continuing evenly about the rounded 
anterior corners into the sides. Posterior angles subrectangu- 
lar. Lateral margin in front of posterior corner with two or 
three weak and wide crenulations. Entire dorsal surface 
densely granular, but with no large tubercles. A little behind 
the anterior border along edge of the darker area a transverse 
row of four widely separated setae. (Pl. 20, Fig. 136.) 

Second tergite a little wider than the collum, its keels bent 
forward, decidedly longer than median portion of tergite, the 
lateral margin of each with an acute tooth at anterior corner 
followed by two crenulations, the posterior corner being a little 
acute and distally rounded. (See Pl. 20, Fig. 136.) The 
third tergite similar to second, but median region a little 
longer actually and relatively to the keels. Fourth tergite with 
keels less bent forward and length of median region obviously 
greater relatively and actually, the four marginal teeth of 
keels the same. Each keel of the fifth tergite has an anterior 
laterally bidentate portion separated by a cleft from an equally 
long projection bearing the pore in its lateral margin, this 
prominence smooth excepting for a few slight granules. The 
other poriferous keels are similar, excepting that three teeth 
are normally evident on anterior lobe. The non-poriferous 
keels have four marginal teeth, these less rounded, more serri- 
form than in anterior segments, with the posterior angle : 


Occasional Papers of the Museum of Zoology 55 


becoming larger and projecting more and more caudad. The 
pore-bearing prominence also comes to project caudad in the 
more posterior segments. (See Pl. 20, Figs. 137 and 138.) 

Dorsal surface of metazonites in general but weakly con- 
vex, densely granular, a transverse furrow but weakly devel- 
oped, two rows of a few more prominent, apically darker, 
granules behind this furrow. Sides of metazonites more finely 
granular. Sterna with granules still finer, largely obsolete, 
and sparer, in part becoming short setae. Spines at bases of 
legs all very short. (See also Pl. 20, Fig. 139.) 

Cauda proper with two setae on rounded tubercles on each 
lateral margin and a pair on dorsum at base. (PI. 20, Fig. 
138.) Anal scale with caudal margin convexly rounded, a 
long seta on each side. (Pl. 21, Fig. 140.) Anal valves 
mesally margined. 

Legs with proximal joints granular. Last joint propor- 
tionately very long. Claws well developed. (For proportions, 
etc., see Pl. 21, Fig. 141.) 

Length, about 30 mm.; width, 4.2 mm. 

Colombia: San Lorenzo. One female taken under a stone 
in a damp creek bed at 4,500 feet; July 3, 1913. Holotype, 
mC. Z.,. 5,144. 


Colombodesmus lygrus, sp. noy. 
Pl, 21, Figs. 142-144 

This is a darker form than the preceding one, the general 
color of dorsum and sides being a dark or dusky brownish 
grey, the poriferous processes alone being lighter, fulvous. 
Venter paler. Legs proximally white or fulvous white, dis- 
tally brownish, the light color extending typically to or a little 
beyond middle of femur. Antennae dark excepting at tip. 

The keels in general form and dentitions similar to those 


56 University of Michigan 


of catharus, but with the poriferous process proportionately 
much smaller and less sharply set off ; this process shortly sub- 
cylindrical, with distal end oblique and having a semicircle of 
granules above and overhanging the pore. (See PI. 2t, Figs. 
142 and 144.) The dorsal surface of metatergite more 
strongly granular; with two transverse series of slightly 
larger, setigerous granules or tubercles behind transverse sul- 
cus and one in front of it, the latter of fewer granules. Sides 
of segment also obviously more concavely and densely granu- 
lar than in the other species. Anal valves and scale more 
strongly granular than in catharus. 

Spines of sterna nearly as in the genotype, but the poste- 
rior pair of each segment larger proportionately to the anterior 
pair. 

For form of leg, see Pl. 21, Fig. 143. 

Length, about 35 mm.; width, 4 mm. 

Colombia: San Lorenzo. One female under leaves at 4,500 
feet; July 21, 1913. Also one female under log at edge of 
clearing at 4,500 feet; July 9, 1913. 

One male, in which the gonopods had unfortunately been 
broken off and lost, under leaves at 2,000 feet; July 16, 1913; 
F. M. Gaige. 

Holotype, M. C. Z., 5,145. 


Cormodesmus, gen. nov. 


Sterna narrower than in Colombodesmus, Trachelodesmus, 
etc. Without sternal spines, but with a rounded tubercle at 
base of each leg on its anteromesal side on all but anterior 
segments. Metatergites wider comparatively between anterior 
corners than between posterior. Keels all well developed, 
thick, laterally lobed through the development of large tuber- 


cles, elevated above dorsum. No repugnatorial pores obvious, 


Occasional Papers of the Museum of Zoology 57 


as in Batodesmus. Metatergites coarsely tubercular, the cauda 
tubercular. Legs rather long, granular. Gonopods of male 
deeply bifid into a broad anterior plate and a posterior branc! 
lying against the former, the posterior branch, through which 


the seminiferous tube runs, with a spur toward its base. 
Genotype, C. hirrutellus, sp. nov. 
In the concealed or absent repugnatorial pores suggesting 
Batodesmus, but readily distinguished from that genus in 


lacking spines on the sterna, in the tubercles of tergites, etc. 


Cormodesmus hirrutellus, sp. nov. 


Pl. 21, Figs. 145-148; Pl. 22, Fig. 149 


The types at present are dark brown, inclusive of legs and 
antennae, excepting that the prozonites may be paler, especially 
below, the color due mostly to adherent material and growths, 
beneath which the color is fulvous. 

Surface of head granular. A series of longitudinal striae 
across base of vertex. Vertigial sulcus and furrow distinct 
to level of antennae. Antennal sockets separated by not more 
than the diameter of the latter. Antennae short, when 
extended straight back not quite attaining caudal edge of col- 
maa. (Pl: ar, Fig: 146.) 


Collum with anterior and lateral margins together forming 
an evenly convex line, excepting for a tubercular projection 
at anterolateral corner below; the posterior corners more nar- 
rowly rounded. Posterior margin weakly convex excepting 
at ends, where more strongly convex. On dorsal surface a 
row of tubercles entirely across anterior border; at each end 
the plate presents an elevation formed by tubercles of the 
anterior series and two caudad of it, the latter series not 


extending across middle region of plate. On the median part 


58 University of Michigan 


of the posterior border two tubercles. Setae as on other ter- 
Sites. (Pl 21; Pig. 145.) 

Second tergite much wider than the collum, its ends or 
keels bent forward, the anterolateral corners appearing as 
thick, rounded tubercles ; the third tergite also bent forward at 
ends. (Pl. 21, Fig. 145.) The following are, as wholes, not 
bent forward at ends, but with the anterolateral corners 
strongly produced forward and similarly rounded, these ante- 
rior processes becoming shorter in going caudad, but project- 
ing as rounded tubercles on all but the last few. The poste- 
rior angles on all but the most anterior keels also come to 
project similarly caudad. Between these two projecting end 
tubercles on a typical segment are two intervening large mar- 
ginal tubercles with a large tubercle below the more anterior 
of them, giving the keel a thick, heavy appearance. Between 
these large marginal tubercles there are typically four, or 
indistinctly five, transverse rows of similar but somewhat 
smaller tubercles, the most anterior row having but two tuber- 
cles, or, if a fifth row is present, this also with two, this form- 
ing the second row, the other usually with six each, inclusive 
of those on keels within marginal ones. Each tubercle bears 
a large seta and numerous very short, fine setae. (See, fur- 
thet, Piyar, Mic 1475) 

Anal valves granulotubercular; mesal borders margined. 
Anal scale trapeziform, the caudal margin straight, with a 
setigerous tubercle at each outer angle. (PI. 21, Fig. 148.) 

Gonopods of male as shown in Pl. 22, Fig. 1409. 

Length, near 35 mm.; width, 5.2 mm. 

Colombia: San Lorenzo. A male and two females from 
stumps at 4,500 feet; July 5, 1913. 

Also two smaller specimens at 5,300 feet; July 23, 1913. 

A not quite adult male in forest at 4,000 feet; July 14, 1913. 


Occasional Papers of the Museum of Zoology 59 


Holotype, M. C. Z., 5,147. 

The dark color of these specimens is due in considerable 
part to foreign material and growths held by the granules and 
numerous short setae, the parts more free from this showing 


a tulvous background. 


Alassodesmus, gen. nov. 


A genus close to Trachelodesmus, but differing from the 
latter in lacking repugnatorial pores on segments 12 and 15, 
and in their presence on segment 14, the pore-formula being 
—, —-, 9, 10, 13, 14, 16-19. Pores large and conspicuous. 


Genotype, A. reductus, sp. nov. 


Alassodesmus reductus, sp. nov. 


Pl. 22, Figs. 150-153 


General color light brown. Legs lighter, more fulvous. 

Differing conspicuously from Trachelodesmus, as repre- 
sented, e. g., by T. ancylophor, in the weaker development of 
the keels, the tergites at the sides merely bulging convexly, 
and the absence of definite polygonal areas above and of the 
larger tubercles, the marginal tubercles being small like the 
others, or a few but slightly larger and more conical. The 
repugnatorial pores open laterally, not elevated on tubercles, 
each surrounded by a rim which may be slightly raised; above 
each pore usually two or three small tubercles, of which the 
caudal one is larger, and on most caudal segments corners to 
project as a small, distally rounded, caudal process. Pores 
present on segments —, 9, 10, 13, 14, 16-19. Dorsal surface 
of metazonites finely roughened, granular or sharpened both 
above and below. Sides of metazonites coarsely granular 
under the keels, the granules becoming finer and sparser below. 
(Pl. 22, Fig. 150.) 


60 University of Michigan 


Sterna not -granular. Spines of sternites of nineteenth 
segment as stout at base as the adjacent joint of the leg, nar- 
rowing distad, curving dorsad, as long as the two first joints 
of leg taken together. The other spines abruptly smaller, 
present forward to the tenth segment inclusive. (See Pl. 22, 
Fig. 751.) 

Cauda short and broad, distally truncate or incurved 
between the two lateral tubercles Anal scale shown in PI. 
22, Fig. 152. 

Legs of same general structure as in Trachelodesmus, with 
claw reduced, etc. Not granular. (Pl. 22, Fig. 153.) 

Width, 3.2 mm. 

Colombia: San Lorenzo. One specimen, unfortunately 
lacking the head and first seven segments, taken in a log at 
about 6,200 feet, July 19, 1913, along with the type of T7ra- 
chelodesmus ancylophor, sp. nov. Holotype, M. C. Z., 5,151. 


Trichomorpha tuberculosa, sp. nov. 
Pl. 22, Figs. 154-156; Pl. 23, Figs. 157-158 
Dorsum black with the keels yellow. .Lower border of 
clypeus yellow. Antennae brown. Legs brownish yellow. 
Dorsum only slightly arched, nearly flat, the keels being 
at a high level. All keels in the male well developed, with 
angles on all from the second caudad acutely and strongly 
produced, becoming particularly long in the caudal region, 
those of the seventeenth to nineteenth segments with distal 
ends distinctly curving mesocaudad. Anterior corners of 
keels rounded, with the usual tooth at lower end of corner, 
and back of this the ectal edge weakly serrate or serratocrenu- 
late, the serrations absent on most caudal keels. Caudal mar- 
gin not truly toothed, bearing usually, however, two setiger- 


ous tubercles. Dorsal surface of prozonites smooth; dorsal 


Occasional Papers of the Museum of Zoology 61 


surface of metazonites strongly tuberculate, the tubercles in 
irregular in part confined transverse series, of which there 
are usually about four caudad of the transverse sulcus, which 
is distinct, and six in front of it, the tubercles in general 
smaller toward the anterior and posterior margins, the tuber- 
cles extending out upon the keels, but just within the margin 
of latter fewer and more weakly developed. All tubercles 
setigerous. (Pl. 22, Figs. 154, 155.) 

Sternites of fifth and sixth segments in the male each with 
a pair of low and broad, distally rounded, large tubercles 
between first pair of legs. 

In the male metatarsal pads projecting distad between the 
tarsi are developed on the first seven pairs of legs. In the 
seventh legs of the male the femur is much thickened, widest 
distad, and is strongly bent, its base projecting as a prominent 
lobe, as shown in Pl. 22, Fig. 156. 

Gonopods small, their form as shown in PI. 23, Figs. 157, 
158. 

Length, 33 mm.; width, 4.6 mm. 

Colombia: San Lorenzo. One male from stump in woods 
at 4,500 feet; July 5, 1913. 

One female from log at 6,200 feet; July 19, 1913. 

Holotype, M. C. Z., 5,152. 


Trichomorpha rugosella, sp. nov. 


Pl. 23, Figs. 159-162; Pl. 24, Figs. 163-164 


Entire dorsum black, excepting the caudolateral corners 
of the keels, which are yellow. Venter brown. Antennae 
light brown. Legs with first joint brown, the second whitish ; 
the third joint or femur and the tibia brown excepting for 


light ring at distal end; last two joints brown or also a little 


62 University of Michigan 


paler at distal end. The legs thus have a more or less banded 
appearance. 

Dorsum but weakly arched, with the keels high and form- 
ing an obtuse angle with adjacent part of metazonites. All 
keels from second caudad with posterior angles acutely and 
strongly produced, the processes becoming longer and also 
curved distally, more curved caudad, though the curvature is 
never strong. Lateral margins of keels nearly smooth, show- 
ing, however, a weak and often vague serration behind the 
anterior corner and usually two similar ones caudad of this 
at points of insertion of setae. Caudal margin of keels not 
dentate (Cf. Pl. 24, Fig. 164.) 

Surface of metazonites with transverse sulcus distinct. 
Both in front and caudad of sulcus the surface is entirely 
divided by deep impressions into tubercle-like elevations which 
are mostly longer than wide and each of which bears a seta; 
there are two series of these areas behind the sulcus and 
three somewhat irregular ones in front, the tubercles of the 
latter more weakly developed in the middle region. General 
dorsal surface of keels convexly elevated, the tubercles over 
this elevation small. (Cf. Pl. 24, Fig. 164.) 

Anal scale as shown in Pl. 23, Fig. 159. 

In the male the tarsal pads of legs cease abruptly at the 
tenth pair. Femur of seventh legs curved, apppearing exca- 
vated below, the proximal end below bulging roundly but with 
no true process. (PI. 24, Fig. 163.) 

Gonopods of male as shown in Pl. 23, Figs. 161-162. 

Length of male, 20 mm.; width, 2.4 mm. 

Colombia: San Lorenzo. A male and female taken in 


forest at 4,000 feet; July 14, 1913; F. M. Gaige. 


Occasional Papers of the Museum of Zoology 63 


A female apparently not in full color and an immature indi- 
vidual probably pertaining to this species were taken at 3,400 
feet, under leaves, on July 15, 1913. 

Holotype, M:. C. Z., 5,154. 


Tricomorpha setosior, sp. nov. 


Pl. 24, Figs. 165, 166 


The type, which may not be in full color, is at present 
-light horn-brown above with the keels light. Sides, venter, 
and legs fulvous. 


This species seems easily distinguishable from the others 
here described by the peculiarities of keels and dorsal sculp- 
turing. The keels have the lateral margin finely but distinctly 
serrate, there being typically on a median segment four to six 
serrations caudad of the usual one back of anterior corner, 
each bearing a seta. The posterior processes of keels in mid- 
dle and posterior regions have the tip more or less bent mesad, 
this curvature being particularly noticeable on the most caudal 
ones. (See Pl. 24, Figs. 165 and 166.) 

All metatergites are strongly densely tubercular, the tuber- 
cles becoming weaker on keels. Tubercles shorter, more 
nearly circular in outline than, e. g., in tuberculosa and rugo- 
sella, and correspondingly more numerous. Three somewhat 
irregular rows back of transverse furrow in addition to a 
marginal series of much smaller tubercles on caudal edge, 
and four or five rows in front of the furrow. Tubercles all 
bearing rather long setae, which form a conspicuous feature. 


Length, about 20 mm.; width, 3.2 mm. 


Colombia: San Lorenzo. One female taken under leaves 
in forest at 4,500 feet on July 20, 1913; F. M. Gaige. Holo- 
mepe, MC. Z.,.5,157- 


64 University of Michigan 


Trichomorpha eutyla, sp. nov. 
Pl. 24, Figs. 167-170; Pi 25, Fig. 171 

Dorsum black, with the keels ferruginous yellow to yellow. 
Antennae dark brown. Legs paler, yellowish brown. 

Dorsum but weakly arched and the keels at a high level. 
Posterior angles of all keels from the second caudad acutely 
and strongly produced. Lateral margins of keels wholly 
smooth, excepting the usual tooth or serration toward the 
anterior corner, this small, the second, third and fourth keels 
also with two weak indentations of serrations caudad of this. 
Caudal margin of all keels caudad of the fifth with a single 
distinct tooth as shown in Pl. 24, Figs. 167 and 168. Processes 
of keels all straight, none at all distally curved. 

Dorsal surface of metazonites smooth, no tubercles at all 
developed ; typically a distinct transverse furrow, from which 
a number of straight, parallel sulci extend caudad, partly 
dividing the caudal region into separate areas. Dorsal sur- 
face of keels moderately elevated and wholly smooth. 

In the legs of the male the pads beneath tarsi are large 
and are strongly and equally developed to the twenty-fourth 
legs, caudad of which they abruptly cease or show as but a 
slight angle over the twenty-fifth legs. (Pl. 25, Fig. 171.) 
Anterior legs and sterna without special lobes or processes. 

Gonopods of male as shown in Pl. 24, Figs. 169, 170. 

Length of male, 20 mm.; width, 2 mm. 

Width of female, to 2.6 mm. 

Colombia: San Lorenzo. Seven males and females taken 
under logs and stones at the edge of a clearing at 4,500 feet; 
July 9, 1913; M. A. Carriker. 

Cincinnati Coffee Plantation, in rotten log; July 2, 1913; 
A. S. Pearse. Several broken specimens, male and female. 

Holotype, M. C. Z., 5,158. 


Occasional Papers of the Museum of Zoology 65 


Trichomorpha eusema, sp. nov. 
Pl. 25, Figs. 174-178 

A species characteristically marked in having the keels and 
the metazonite of each ordinary segment entirely fulvous 
caudad of the transverse sulcus and in a median spot in front 
of it, the anterolateral portion of metazonite within the keel 
and the prozonite deep chocolate brown to nearly Dlack. 
Antennae light brown. Legs fulvous. 

Dorsum low, with keels high and elevated. Posterior 
processes of keels strongly developed, as usual, straight or 
only slightly curved at tip in a few of the most caudal. Lat- 
eral margin of keel with a distinct serration caudad of the 
rounded anterior corner, otherwise smooth. A small, acute 
tooth at angle between posterior process and caudal margin 
of metazonite proper. (See Pl. 25, Figs. 174 and 175.) 

Tergites above with a deep transverse sulcus; area behind 
this sulcus, divided by longitudinal sulci into longitudinally 
oblong elevated areas extending to caudal margin or with pos- 
terior ends set off as smaller setigerous tubercles, forming a 
second row along the margin, typically ten or eleven areas in 
each series. Region in front of transverse furrow with no 
such areas set off by sulci, smooth or nearly so. 

Seventh leg of male shown in Pl. 25, Fig. 176. 

Gonopods of male as shown in Pl. 25, Figs. 177, 178. 

Length, 24 mm.; width, 2.6 mm. 

Colombia: San Lorenzo. Two females taken in forest at 
4,000 feet; July 14, 1913; F. M. Gaige. 

One female under a stone in a damp creek bed at 4,500 feet , 
July 3, 1913. ° 

One male and one female from stump in forest at south 
end of Cincinnati Coffee Plantation ; July 5, 1913; F. M. Gaige: 

Holotype, M. C. Z., 5,161. 


66 University of Michigan 


Trichomorpha angulella, sp. nov. 


Pl25.. Bas: 75 


General color of dorsum and keels brown, in part of a 
chestnut cast, excepting only the caudal processes of the keels, 
which are fulvous. Antennae brown. Legs fulvous. 

Convexity of dorsum and elevation of keels as usual. Pos- 
terior processes of keels straight, more than usually slender 
and acute, especially on the more posterior segments. Lateral 
margin of keels wholly smooth excepting the usual serration 
caudad of the anterior corner, this distinct. Caudal margin 
wholly untoothed and smooth. (See Pl. 25, Fig. 173.) Meta- 
tergite with transverse sulcus distinct. 

Surface in front of this sulcus wholly smooth and bearing 
a transverse series of widely separated setae; surface caudad 
of the sulcus a little roughened by rather fine longitudinal or 
in part somewhat oblique impressed lines; a series of setae 
along caudal border and one farther forward, the latter typ- 
ically of six setae in all. 

Length, about 16 mm. ; width, 1.6 mm. 

Colombia: San Lorenzo. Two adult and one immature 
female at 2,500 feet; July 15, 1913. Holotype, M. C. Z., 5,165. 

Trichomorpha paurothrix, sp. nov. 


Pl. 25° Fic. 172 


Dorsum black, excepting the outer border and posterior 
processes of keels, which are yellow; the mesal, convex area 
of keel mostly dark like the dorsum in general; the processes 
of keels bright yellow, the anterior light portion of a more 
vague cast. Antennae dark brown, the legs a lighter, fulvous 


brown. 


Occasional Papers of the Museum of Zoology 67 


Dorsum and relation of keels as usual. Keels with lateral 
margin presenting the usual serration behind rounded anterior 
corner, otherwise smooth, a single seta caudad of the serra- 
tion; caudal margin not toothed. (See, further, Pl. 25, Fig. 
fg2;) 

Metazonites with transverse sulcus deep, a median longi- 
tudinal sulcus also distinct. The region in front of the sulcus 
is further divided on each side by a weaker longitudinal fur- 
row, thus giving a larger mesal area and a smaller one adja- 
cent to the keel. The region behind the transverse sulcus with 
two large, rounded areas on each side adjacent to the sulcus 
and four much smaller areas or tubercles on each side along 
caudal border, these decreasing in size from the outer to the 
mesal end of the series. Each of the larger anterior areas 
bears a seta; but the surface of the metazonites is otherwise 
glabrous. Dorsal surface of keel moderately convexly ele- 


vated, the surface smooth. 
Length, about 24 mm.; width, 3 mm. 


Colombia: San Lorenzo. July 22, 1913; one female from 


a bromeliad on tree at 5,000 feet. Holotype, M. C. Z., 5,167. 


Chondrodesmus tamocalanus, sp. nov. 


Pl. 26, Fig. 179-182 


Dorsum black or nearly so, a narrow stripe along caudal 
border of metazonites sometimes lighter, reddish, and the keels 
yellow, the line limiting the yellow area running obliquely 
from the mesal end of caudal edge of keel to cephalolateral 
corner, the cephalolateral region commonly tinged with red- 
dish. Cauda yellow. Collum narrowly bordered in front and 


behind with yellow and its keels entirely yellow. Antennae 


68 University of Michigan 


and legs of a pronounced pink tinge, this color deeper distad, 
the proximal joints showing some fulvous. 

Dorsum but moderately arched, the keels horizontal. 

The collum moderately narrowed down the sides. Ante- 
rior margin curving widely over anterolateral corner around 
to caudolateral corner, which in its general outline is subrec- 
tangular but with apex rounded. Surface smooth, wholly 
lacking granules, under the lens at most showing a network 
of fine coriarious lines. 

On the metazonites in general tubercles wholly. absent or 
at most only obscurely indicated in part. The entire surface 
with network of deep coriarious impressed lines. 

Keels of segments two to four with anterior corners 
rounded, the posterior rectangular and the lateral margin 
straight, the margins wholly smooth. On the sixth segment 
the caudal of keels has become acute, the production of the 
angle gradually increasing on successive segments in going 
caudad. Anterior and lateral margins of keels smooth and 
even throughout. Caudal margin mostly with a single, low, 
obtuse tooth at middle, or else wholly smooth. (PL. 26, Figs. 
179-181. ) 

Gonopods of male as shown in Pl. 26, Fig. 182. 

Length of male type, 38 mm.; width, 7.6 mm. 

Width of female, 8.8 mm. 

Colombia: Near Mamatoca, at foot of mountain; July 28, 
1913. A male and two females taken under leaves along a 
ditch from the Tamocal River, elevation 200 feet. 

Fundacion River; Aug. 12, 1913. A recently moulted male 
taken under log. It is now light brown-in color, with the legs 
reddish in typical manner. It is 9 mm. long. 


Cincinnati Coffee Plantation; July 5, 1913; F. M. Gaige. 


Occasional Papers of the Museum of Zooloyy 69 


One recently moulted female agreeing in appearance with the 
preceding male. | 

San Lorenzo; July 1, 1913. One male on trail between 
2,000 and 4,500 feet, in heavy forest. 

Fioterype, WS ©? 25 °5, 168. 

This species in the exceptionally small size and general 
structure of the male gonopods is obviously close to the Ecua- 
dorean C. armatus Silvestri and the Colombian C. riparius 
Carl. The terminal process of the gonopods is very similar 
to that of riparius, though the sickle-shaped end portion is 
somewhat less slender and the broad tooth proximad of it 
arises from the ectal edge, not from the mesal, as described 
and figured by Carl for riparius. It has a tooth on the caudal 
margin of the keels not present in riparius. The keels are 
yellow forward to anterior corner instead of only back of 
middle. 


Chondrodesmus cerasinopus, sp. nov. 


Pl. 26, Figs. 183, 184 


A species resembling the preceding one. The dorsum deep 
brown rather than black, with the light color of keels not so 
bright a yellow, more horn-colored, and continuing farther 
forward on the anterior corners. The entire head is dark, 
brownish black, the clypeal region not pale as it is in the other 
species. Legs and antennae contrasting in being of a uniform 
dark cherry red. 

The dorsum is more strongly arched and the keels more 
depressed. The posterior angles of keels in general less pro- 
duced and lacking an obtuse tooth on posterior margin, this 
being often finely granular or uneven. The marginal thicken- 


ing of the keel on the poriferous segments is notably more 


70 University of Michigan 


elevated, with mesal face of elevation commonly projecting 
over base, grooved. (PI. 26, Figs. 183, 184.) 

The surface of the metazonites is more strongly rough- 
ened, with distinct areas in considerable part developed and 
a transverse furrow. more evident on some of the segments. 

Length, 60 mm.; width, 10 mm. 

Colombia: Fundacion River; Aug. 19, 1913. One female 
taken under a log. Holotype, M. C. Z., 5,173. 


Chondrodesmus virgatus, sp. nov. 
Pl. 26, Figs. 185, 186 


The animal is conspicuously marked dorsally with alter- 
nating cross bands of dull reddish brown and yellow, con- 
siderably more than the caudal half of each metazonite being 
yellow, the remaining portion and the prozonite being of the 
darker color. Cauda proper yellow. Collum with yellow 
caudal border and anterior border, which does not extend to 
ends. Venter yellow. Head uniformly reddish. Legs red- 
dish excepting proximally. Antennae yellow distally, proxi- 
mally with reddish tinge. 

Collum strongly narrowed at sides in the usual manner. 
Surface wholly smooth. 

All keels narrow. Tergites two to four with outer por- 
tion narrow ectad, both corners of keel well rounded and 
obtuse in general outline. Posterior angle of the keels of 
the sixth segment is the first to be a little produced. Posterior 
margins of all keels smooth and even, or at most very minutely 
and irregularly uneven. All poriferous swellings are sharply 
set off from the ordinary marginal rim in front, each lanceo- 
late in outline with apex cephalad. See, further, Pl. 26, Figs. 
185 and 186. Surface without transverse sulci and otherwise 


wholly smooth. 


Occasional Papers of the Museum of Zoology re 


Length, about 35 mm.; width, 6.5 mm. 

Colombia: Fundacion; Aug. 19, 1913. One adult female 
and one immature specimen taken under log on Fundacion 
River. Also one female in same locality taken Aug. 12, 1913. 
Holotype, M. C. Z., 5,174. 


Chondrodesmus virgatus, var. frater, var. nov. 


Similar in appearance to the preceding species, but the yel- 
low color covering the entire metazonite on all segments, except- 
ing a narrow stripe along anterior margin between the kecls, 
which is more or less suffused with the reddish color of the 
prozonite. Keels entirely yellow. Collum with median region 
reddish, the border yellow all the way around, the anterior 
yellow stripe not restricted to median region as it is in wir- 
gatus. Venter yellow. Antennae wholly yellow. Legs mostly 
yellow, a reddish tinge weak and uneven in some. 

Surface of metazonites wholly smooth as in wirgatus. 
Form of keels in general similar; but in the present form the 
posterior angle of the fifth keels is distinctly produced, not 
at all rounded as in virgatus. The posterior angle of the sey- 
enteenth keel is also more produced. 

Length, about 40 mm.; width, 6.5 mm. 


Colombia: San Lorenzo; July 25, 1913. One female taken 
under a log at 2,000 feet. Holotype, M. C. Z., 5,177. 


Chondrodesmus rugosior, sp. nov. 


Pl. 27, Figs. 187, 188 


Dorsum brown, with keels pale, light olive grey to olive 
yellow, over entire length. Collum with only the keels light, 
there being in the type no pale anterior and posterior borders. 


Head brown, not paler in clypeal region. Legs at present 


72 University of Michigan 


dilute olive yellow to olive grey, the distal articles having a 
weak pink tinge. All articles of antennae with pink tinge. 
Margin of collum curving widely and evenly about each 
antero-lateral corner to caudal corner, which is nearly rectan- 
gular in general outline, though with its apex founded as 
usual. Surface smooth excepting for the usual network of 


fine impressed lines. 


The anastomosing lines of the metazonites of the segments 
of median and posterior regions much more deeply impressed, 
producing a strong roughening which is heightened by pro- 
nounced rugae, particularly on the more posterior plates; dis- 
tinct polygonal areas are also set off. Low tubercles irregu- 
larly developed, particularly along caudal borders. Keels mod- 
erately broad. Pore swelling evenly passing into ordinary 
marginal elevation in front, not set off. Posterior angles of 
keels not truly produced in segments in front of the sixteenth, 
the angle bulging slightly caudad first on the seventh keels. 
Posterior margin of keels often rather finely uneven, but not 
presenting a true tooth, or such only occasionally indicated. 
See further Pl. 27, Fig. 188. 

In the gonopods the terminal blade is much broader than 
in those of tamocalanus, etc., and the major tooth below the 
apical prong is on the mesal edge as in riparius Carl, the outer 
edge presenting merely several low serrations. See further 
Pl. 27, Fig. 188. 

Length, about 48 mm.; width, 9.5 mm. 

Colombia: San Lorenzo; July 5, 1913. One male taken 
in a stump in woods at 4,500 feet. Holotype, M. C. Z., 5,178. 


Occasional Papers of the Museum of Zoology a2 


Zigwadesmus guiananus, sp. noy. 
Pl. 27, Figs. 189-193; Pl. 28, Fig. 195 

Color above and over sides brownish chestnut to chocolate, 
the venter more fulvous. Keels and an immediately adjoining 
portion of tergite above them yellow, as is also a narrow 
median dorsal longitudinal stripe which is interrupted on ante- 
rior part of each prozonite. Legs light brown. No pale spot 
.on each side of prozonite such as characterizes brunneus. 
Antennae darker brown. 

Head widely and deeply excavated from base of each 
antenna caudad, an elliptic, somewhat elevated organ lying in 
this excavation, as in the genotype. 

Keels of second, third and fourth segments normally devel- 
oped. Each with a low tooth at anterior corner, which is bet- 
ter developed than in brunneus. Caudal angle of fourth keels 
more broadly produced than in brunneus. Keels caudad of 
the fourth narrow and thick, appearing as simple bulgings of 
the tergite with a margining sulcus; caudal angles produced 
in all, but keels in anterior portion becoming low and obsolete, 
though in general a little better developed than in the genotype 
and inserted a little higher on the sides of the segments. Porif- 
erous process set off by a sharp sulcus from remaining por- 
tion of keel, its ectal face elongate elliptic. (Pl. 27, Figs. 189- 
191.) 

The conical tubercles at distal end of cauda are shorter and 
more rounded than in brunneus and are not directed somewhat 
ventrad of caudad as in that species. (See Pl. 27, Fig. 193, 
and Pl. 28, Fig. 195.) 

The anal scale differs from that of brunneus in having the 
caudal margin between setigerous tubercles rounded, instead 
of angular, and not surpassing the tubercles. (PI. 27, Fig. 


192. ) 


74 University of Michigan 


Length, 38 mm.; width, 5.2 mm. 
British Guiana: Cacao Plantation about camp. One 
female taken in rotten wood, July 18, 1914; F. M. Gaige. 


Also one female in “Forested Sand Hills,” Aug. 18, 1914; 
F. M. Gaige. 


Holotype, M. C. Z., 5,170. 
This and the following one are the first species to be 


referred to this genus in addition to the genotype, Z. brunneus, 


the latter having been described from Trinidad. 


Zigwadesmus modestus, sp. nov. 


Pl. 27, Fig. 194; Pl. 28, Figs. 196-199 


Brown, the keels and a small area just above each paler, 
as usual. No middorsal stripe and no light spots on sides of 
prozonites. 


The anterior keels as in the other species, the angulation 
of the fourth more as in brunneus. ‘The posterior angles of 
keels in general less produced and less acute than in the other 
two known species. The poriferous processes are obviously 
different in being directed more nearly ectad, with caudal angle 
more rounded; section in the anterocaudal direction shorter, 
the whole more approaching a cylindrical form. (See Pl. 28, 
Figs. 196, 197, and 198.) 

The cauda much as in guiananus, but the terminal tubercles 
or processes even shorter. (Pl. 28, Fig. 199.) Anal scale 
with caudal margin between setigerous tubercles obtusely angu- 
lar, extending beyond the, tubercles. (Pl. 27, Fig. 194.) 


Length, about 22 mm.; width, 3.25 mm. 


British Guiana: Forested Sand Hills. One female of 
nineteen segments taken in sandy’ soil of forest floor. Aug. 
14, 1914; F. M. Gaige.. Holotype, M. C. Z., 5,181. 


Occasional Papers of the Museum of Zoology 75 


CRYPTODES MIDAE 


Arionus, gen. nov. 


Body composed of head and twenty segments. 


Collum covering the head; its anterior border divided into 
ten areas by radial lines as in Docodesmus, etc., elsewhere 
granular. | 


Second tergite broader than collum and than third tergite, 
its lateral border showing these areas separated by short radial 
lines. Other tergites with keels comparatively narrow; pre- 
senting three lateral areas separated by impressed radial lines 
and slight marginal indentations, or occasionally on one or 
both sides, in more posterior segments four lobes. Caudal 
margin of keels with one incision. Keels continuing nearly 
the general outline of dorsum. Caudal border of metatergites 
divided into a series of areas by longitudinal impressions, the 
margin correspondingly weakly crenate. Pores inconspicuous, 
on caudal lobe back from margin; occurring on segments five, 
seven, nine, ten, twelve, thirteen, fifteen to ‘nineteen. Dorsal 
surface of metatergites coarsely granular or tubercular, tuber- 


cles nearly uniform. 

Penult tergite with caudal margin convex, the areas, of 
which there are six in all in the genotype, produced caudad as 
stout acute teeth, of which the lateral one on each side or 
process of keel is not larger than others, which accordingly 
extend farther caudad. 


Anal tergite!small, now lobate, with cauda exposed from 
above. 


Gonopods of male with coxae large. ‘The telopodite also 


large and stout, exposed in lateral view. 


Genotype, A. uwlophilus, sp. nov. 


76 University of Michigan 


Arionus ulophilus, sp. nov. 


Pl. 28, Figs. 200, 201; Pl. 29, Figs. 2c2 2c6 


Dorsum (metazonites) brown, paler in going caudad. Ven- 
ter paler, fulvous. 

Vertex of head covered by an elevated shield-shaped and 
strongly tubercular area, ‘this area crossed by a longitudinal 
sulcus ending in a notch in the lower end of the elevated area. 
This area brown, the lower smoother parts of head fulvous. 
Face with a large, rounded prominence on each side ectad of 
antennal socket. (Pl. 29, Fig. 202.) ; 

Collum with anterior margin convex, the border in form 
of a rim divided by sulci and slightly marginal crenations into 
ten lobes. ‘The surface otherwise densely tuberculate, all of 
the tubercles small. (Pl. 29, Figs. 202, 203.) 

Second tergite bowed forward at ends so that basal lobes 
of posterior margin extends ectad and appears as a shorter 
fourth lobe. This and following segments show an anterior 
rim divided into areas similar to those of caudal border, these 
becoming shorter in going caudad. (PI. 29, Fig. 203.) Dor- 
sum of metazonites densely tubercular, none of tubercles large, 
forming five or six more or less irregular transverse rows. 
Pores small, scarcely elevated. (PI. 28, Fig. 201.) 

On antepenult tergite the caudal areas project as obtuse 
processes with the caudal processes of keels large; these proc- 
esses on the penult are longer, more acute and six in number 
inclusive of the processes of keels which are of similar size. 
(Pl. 29, Fig. 204.) 

Anal tergite as shown in Pl. 29, Fig. 204. Anal scale, PI. 
28, Fig. 200. 

Gonopods of male as represented in Pl. 29, Figs. 205 and 
200. 


Occasional Papers of the Museum of Zoology a7 


Length, 11 mm.; width, 1.9 mm. 
Colombia: San Lorenzo. One male in forest at 4,000 feet, 
July 14, 1913; F. M. Gaige. Holotype, M. C. Z., 5,182. 


Guianonus, gen. nov. 


A genus resembling the preceding one, Arionus, in having 
the keels narrow and in having the metatergites densely cov- 
ered with numerous coarse granules or tubercles, of which 
none are specially enlarged. ‘The keels are more elevated thar 
in the other genus, standing out nearly horizontally. Keels. 
laterally fundamentally three-lobed excepting most posterior, 
which are four-lobed. Posterior processes of keels of penult 
tergite strongly developed, clearly surpassing the median por- 
tion of caudal margin. At once distinguishable from Arionus 
in the position and distribution of the repugnatorial pores, 
which open at the distal end of the caudal lobe of keels of 
segments five, seven, nine, ten, twelve, thirteen, fifteen, and 
sixteen. Cauda simple, exposed from above. 

Telopodite of gonopods of male large and exposed. 


Genotype, Guianonus ectoporus, sp. nov. 


Guianonus ectoporus, sp. nov. 


Pl. 29, Figs. 207-209; Pl. 30, Figs. 210, 211 


Dorsum dusky brown or chestnut, collum and head lighter, 
more ferruginous. Venter fulvous brown. 

Vertex of head somewhat raised as in Arionus; but the 
area only obscurely tubercular above and below, more distinctly 
so across its median region. 

Collum with anterior rim presenting twelve areas, the one 
at each end of the series smaller than the others. 

Second tergite considerably wider than collum and than 


following tergites; moderately bowed forward at ends, but 


78 Unizersily of Michigan 


much less strongly so than in, e. g., Arionus ulophilus; keel 
longer than median region of metatergite, its anterior lobe 
large and rounded. (Pl. 30, Fig. 210.) In the third and 
fourth metatergites the keels are not longer than the middle 
region; anterior lobes smaller; anterior margin incised each 
side of a distinct lobe. (Pl. 30, Figs. 210, 211.) Keels of 
other segments shorter than median region of metatergite. On 
the keels of the fifth and succeeding segments the anterior 
lobe becomes smaller and is bent upward, soon appearing 
merely as a tubercle, while the lobe of the caudal margin 
remains distinct and is usually distally obtusely angular. The 
lateral lobes of keels, especially the anterior ones, also tend 
to be bent more or less upward. ‘The porigerous lobe is typic- 
ally bifid at the tip, the pore opening between the two sub- 
divisions a little above the edge, not elevated on a secondary 
cone or eminence. ‘The caudal border of segments divided into 
lobes by longitudinal furrows and corresponding crenations as 
in Arionus, the crenations becoming deeper on the more caudal 
segments. Dorsal surface of segments densely coarsely gran- 
ular, two granules in each of }four longitudinal rows on each 
metatergite a little larger, more tubercle-like than the others, 
these tubercles more distinct on posterior segments. (See also 
Pl. 29, Fig. 207.) Keels of seventeenth to nineteenth ‘seg- 
ments four-lobed and having posterior angles produced, the 
processes on nineteenth segment obviously larger than the 
caudal teeth or crenations of the median region fof segment. 
(See PL 20, Fig. 207.) 

Cauda in general outline posteriorly rounded, but with a 
short distal piece set off and truncate across end; across basal 
region a transverse series of four tubercles, the end one on 
each side projecting as a distinct lateral process or tooth; sur- 


face in front of these tubercles with smaller granules. (PI. 


Occasional Papers of the Museum of Zoology 79 


29, Fig. 207.) Anal valves margined. Anal scale triangular, 
a tubercle on each side of caudal angle. (Pl. 29, Fig. 208.) 

Gonopods of male as shown in PI. 29, Fig. 209. 

Length, 1o mm.; width, 1.3 mm. 

British Guiana: Second Mourie. Aug. 20, 1914; F. M. 
Gaige. Two males collected among the dead leaves on the 
ground in a tree clump. 

Sand Hill Forest, on headwaters of Hubidibu Creek. Aug. 


26, 1914; F. M. Gaige. Three smaller females taken in sandy 
forest floor. 


Holotype, M. C. Z., 5,183. 


CyYRTODES MIDAE 


Cliodesmus, gen. nov. 


Body composed of head and twenty segments. 

Collum narrow, not covering the head above. 

Second tergite much wider, the keel expanded, its anterior 
border carried forward. Caudal margin of keel of second and 
third segments with a projection above a transverse furrow 
into which the edge of the succeeding keel fits. Keels of seg- 
ments in general bent downward at outer ends, the ectal mar- 
gin rounded and crenate. Anterior and posterior margins 
entire. Dorsum strongly arched. Each metatergite with two 
transverse rows of large tubercles, the collum with three; no 
granules or setae. Pores on upper part of anterior slope of 
keels except on segments fifteen to nineteen, where they have 
shifted to the outer slope; present on segments five, seven, nine, 
ten, twelve, thirteen, and fifteen to nineteen. 

Nineteenth segment extending completely over the twen- 
tieth, its keels meeting behind as in Cyrtodesmus, etc. 

Coxae of male gonopods large, concealing the telopodite. 

Genotype, Cliodesmus cryptopygus, sp. nov. 


80 University of Michigan 


Cliodesmus cryptopygus, sp. noy. 


Pl. 30, Figs, 213-217 


Vertex of head and metazonites, together with the normally 
exposed portion of prozonites, black above, the covered por- 
tion of prozonites and the anterior portion of each keel normally 
overlapped by the preceding keel, pale, whitish. Lower part 
of head, antennae, and venter in general with legs pale, fulvous 
or whitish fulvous. 

Head granular in a cross band between and just above 
antennae, elsewhere nearly smooth. A transverse depression 
across clypeus. Antennae with joints short, as a whole first 
clavately widening and then again narrowing at distal end. 
(PI. 30, Fig: 213;) 

Collum with ends or keels rounded, each above with a large, 
transversely elongate ‘or somewhat double tubercle. Median 
region of plate with three transverse rows of large tubercles, 
six in each row, the ectal tubercle of the two anterior rows on 
each side in some degree fused with each other at base of keel. 
(Pl. 30, Fig. 213.) 

Second tergite evenly and moderately bending forward on 
each side; anterior margin entire; lateral margin with five cre- 
nations, of which that at anterior corner is largest; caudal 
margin of keel deeply excavated at base of distal depressed 
portion, leaving an angle mesad of it which projects over a 
transverse furrow in caudal edge of plate into which the ante- 
rior border of the succeeding tergite fits. (Pl. 30, Fig. 213.). 
Dorsum of tergite with two transverse rows of tubercles, six 
in each series, and in addition two partially confluent tubercles 
on keel in a line just mesad of the caudal excavation, the keel 
elsewhere smooth. Third tergite with keels not expanded as 
those of the second are, there being but three lateral crena- 


Occasional Papers of the Museum of Zoology 81 


tions; caudal excavation of keel and the tubercle of segment 
as in the second. In the fourth and succeeding keels there is 
no excavation on the caudal side, the anterior border being 
bent down so as to bend beneath the preceding keel as a whole 
instead of fitting into a furrow and notch. Fourth keels with 
four crenations on lateral margin, the following ones with six. 
Tubercles in general as described for the second segment, 
excepting that typically only one tubercle is distinct at the bend 
in the keel. 

Penult tergite widely overlapping the last segment, the keels 
meeting behind, or separated only by a narrow incision; caudal 
end of each keel rounded, four crenations between the rounded 
caudal end and the widely rounded anterior corner; dorsum 
convexly elevated above the keels, which are pale in color 
throughout, and bearing two transverse rows of tubercles, six 
in each row. The keels curve down marginally all the way 
around, (Pl. 30, Fig. 214.) 

Anal segment with cauda short and rather narrow, distally 
rounded. Valves a little convex, mesally margined. (PI. .30, 
Fig. 215.) Scale subtrapeziform. (See Pl. 30, Fig. 215.) 

Gonopods of male with coxae, completely concealing the 
telopodites when in situ. See further Pl. 30, Figs. 216 and 217. 

Length, 8 mm.; width, 1.2 mm. 

British Guiana: First Mourie. Aug. 26, 1914; F. M. 
Gaige. A male and a female collected under fallen leaves in 


a tree clump. Holotype, M. C. Z., 5,187. 


82 University of Michigan 


Agnurodesmus thrixophor, sp. nov. 


Dorsum brown over fulvous, the brown pigment on each 
metatergite tending to outline roughly two transverse rows of 
polygonal area. Venter and legs fulvous. 

Collum with anterior margin straight; the caudal and lat- 
eral margins together forming a convex or semi-circular curve. 
Across the anterior border a series of longitudinal sulci, leaving 
between them ridges or elongate tubercles, each of which bears 
a seta; this series of tubercles continues out upon the lateral 
lobe or keel, but shifts a little caudad, the tubercles being 
shorter and more remote from the anterior margin. Back of 
this anterior series of ridges the surface bears numerous shorter 
setae, :\(BLy 303 Pig. 212.) 

The second tergite is long, bent downward and expanded 
below, the lower margin attaining the lower level of head and 
the anterior rounded corner protruding forward also as far 
as the anterior edge of the head or nearly so. Lateral margin 
with ten crenations, the first being the much larger rounded 
anterior corner. Caudal margin notched somewhat as in Clio- 
desmus, but the notch larger and more obtuse. Keel margined 
all the way around. 

The third tergite with keel much shorter, only four crena- 
tions on ectal margin. Caudal border concavely excavated for 
reception of succeeding keel. In the following segments the 
anterior margin smooth and straight, or nearly so, the mesal 
angle jutting forward as usual. Four crenations at ectal end 
of fourth to sixth keels, usually five on the others or six on a 
few of the last ones. Within the margining rim all around, 
or more vaguely on anterior border, a series of areas sepa- 


rated by sulci or depressions, each one bearing a seta; the 


Occasional Papers of the Museum of Zoology 83 


margin often vaguely crenate in correspondence with all or 
most of these border areas. The fourth and immediately fol- 
lowing keels narrowing and rounded distad; those of more 
posterior segments more quadrate. (PI. 30, Fig. 212.) Cau- 
dal margin of keels of last few segments oblique, the caudal 
angle becoming acute, those of the eighteenth more strongly 
bent back, while those of the nineteenth nearly meet in the 
middle line and widely overlap the last segment in the usual 
manner. Across each ordinary metatergite two rows of long 
setae, typically six setae in each series between the keels. 

Cauda of anal segment narrow, distally rounded, not reach- 
ing caudal edge of keels of nineteenth segment. Anal scale 
broad, triangular, the caudal angle extending beyond the 
tubercles. 

Length, about 16 mm.; width, 2.5 mm. 

Colombia: San Lorenzo. July 3, 1913; F. M. Gaige. One 
female under log at 4,500 feet. Holotype, M. C. Z., 5,188. 

Easily distinguished from A. verrucosus (Brolemann), the 
genotype, by differences in the collum. In the latter species 
the collum is crossed by two series of setae in which those of 
the posterior series are larger. In thrixophor, on the contrary, 
there is'an anterior series of long setae, while the remaining 


part of the plate bears shorter setae. 


84 


Fig. 


Fig. 


Fig. . 


Fig. 


Vig. 


University of Michigan 


Pirate I 


Glomeridesmus orphnius, sp. nov. 


Collum and tergite. x 45. 


Lower part of tergites XIII, XIV and XV. 


Siphonophora graciliceps, sp. nov. 
Anterior end, dorsal view. 
Anterior pleurite. x75. 
Pleurite of posterior region. x75. 
Siphonophora pearsei, sp. nov. 


Rostrum, etc., in side view, outline. x 30. 


Pleurite of right side, anterior region. x75. 


Pleurite of left side, posterior region. x 76. 


X 45. 


Souru AMERICAN DIPLOPODA PLATE I 


86 University of Michigan 


PiLate II 


Siphonophora pearsei, sp. nov. 


Fig. 9. Anterior end, dorsal view. x 30. 
Fig. 10. Leg of sixth segment of male. x 76. 
Fig. 11. Leg from posterior region of male. x 76. 


Fig. 12. Gonopods of male, posterior view. x7 


on 


Fig. 13. Distal portion of left anterior gonopod, mesal view. 


Fig. 14. Posterior gonopod. x 112. 


Fig. 15. Tip of posterior gonopod. x325. 


Siphonophora corynetes, sp. nov. 


Fig. 16. A pleurite of right side of anterior region. x 76. 


xX TI2Z 


PLatsE II 


SoutH AMERICAN DIPLOPODA 


88 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


Fig. 


17. 


18. 


19. 


20. 


22. 


23. 


University of Michigan 


Prams il 


Anterior end, dorsal view. x 47. 


Pleurite from right side of posterior region. 


Anterior gonopod. x 185. 


Posterior gonopod. x 18s. 


Siphonophora relicta, sp. nov. 


Anterior end, dorsal view. x45. 


Head and collum in outline, from in front. 


Right pleurite from anterior region. 


x75: 


x75: 


x 45. 


SoutH AMERICAN DIPLorova PLate III 


17 18 


ele) 


Fig. 


Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


28. 


20. 


University of Michigan 


PiLate IV 


Siphonophora relicta, sp. nov, 


A left pleurite from posterior region. x75. 


Siphonophora gwianana, sp. nov. 


Anterior end, dorsal view. x 30. 
Left pleurite of anterior region. x 76. 


Left pleurite from posterior region. x75, 


Siphonotus parvus, sp. nov. 


Head and collum from in front. x I12. 


Caudal end, from above. x75. 


SoutH AMERICAN DIPLOPODA PLatTe IV 


30; 


Pegi 


ae 


aa Se 


gers 


University of Michigan 


PLATE V 


Stemmiulus craurus, sp. nov. 


Gnathochilarium of male. x 76. 


Gnathochilarium of female (setae omitted). 


Second legs of male. x45. 
Tip of third leg of male. x 765. 


Gonopods of male, posterior view. 


x 46. 


x75. 


SoutH AMERICAN DIPLOPODA Prt Vv 


9+ 


Fig. 


Fig. 


36. 


aie 


ig. 38. 
ase 
ig. 40. 


g. 41. 


University of Michigan 


Pirate VI 


Stemmiulus craurus, sp. nov. 
First legs of male. x 45. 


Third legs of male. x45. 


Stemmiulus drymophilus, sp. nov. 
Gnathochilarium of male (setae, excepting upper laterals, 
omitted). x45. 
First leg of male. x 45. 
Second legs of male, anterior view. x 45. 
Third leg of male. x 45. 


Tip of third leg of male. x 325. 


South AMERICAN DIPLOPODA Prats VI 


QM; 


96 


Fig. 
Fig. 


Fig. 


Fig. 


Vig. 
Tig. 


Tig. 


42. 
43- 


44. 


45. 


46. 


47. 


48. 


University of Michigan 


Pirate VII 


Stemmiulus drymophilus, sp. 


nov. 


Second leg of male, ectal view. x 46. 


Tip of second leg of male. x 325. 


Gonopods of male, anterior view (setae from distal end proL- 


ably rubbed off). x 30. 


Gonopods of male, posterior view. 


X 30. 


Stemmiulus ruthveni, sp. nov. 


Gnathochilarium of male. x55. 
First legs of male. x 46. 


Second legs of male, posterior view. 


x 46. 


SoutH AMERICAN DIPLOPODA Peate? VII 


98 University of Michigan 


Pirate VIII gh ie 


Stemmiulus ruthveni, sp. nov. 


Fig. 49. Third legs of male. x 45. 
Fig. 50. Tip of third leg of male. x 325. 
Fig. 51. Gonopods of male, posterior view. x 30. 


Fig. 52. Distal ends of telopodite of gonopod, anterior view. x73. 


Stemmiulus labbanus, sp. nov. 


Fig. 53. Gnathochilarium of male. x 45. 
Fig. 54. First leg of male. x 45. zs. j 


Fig. 55. Tip of second leg of male. x 325. 


PLATE VIII 


SoutH AMERICAN DIPLOPODA 


I0O 


Fig. 
Fig. 57. 
Fig. 5 
Fig. 5 


Fig. 


Tig. 


Or. 


University of Michigan 


Prats 1X 


Stemmiulus labbanus, sp. nov. 


Second leg of male. x45. 
Third leg of male, caudal view. x 45. 


Tip of third leg of male. x325. 


Gonopods of male, posterior view. x 30. 


Gonopods of male, anterior view. x 30, 


Prostemmiulus heterops, sp. nov. 


Caudal edge of ventral end of twentieth tergite. 


Soutin AMERICAN DIPLOPODA q PLATE IX 


102 


Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


63. 


64. 


67. 


68. 


University of Michigan 


Prare X 


Prostemmiulus heterops, sp. nov. 


Gnathochilarium of female. x55. 
Head and collum from right side, female. x 30. 


Portion of head of same from left side to show eyes. x 30. 


Epinannolene lorenzonus, sp. nov. 


Collam and adjacent parts, left side. x17. 


Right gonopod, posterior view (setae probably in part rubled 
OiD ets KIO; 


Right gonopod, anteromesal view. x 30. 


Epinannolene artus, sp. nov 


Collum of female. x 45. 


SoutH AMERICAN DIPLopPopA oy! Pirate X 


104 University of Michigan 


Pirate XI 


Epinannolene arius, sp. nov. 


Tig. 69. Gonopods of male, posterior view. x75. 


Epinannolene xestus, sp. nov. 


Fig. 70. Collum and portion of head, lateral view. x 30. 


Typhlonannolene adaptus, sp. nov. 


Fig. 71. Anterior end, lateral view. x 30. 
Fig. 72. Gnathochilarium of female. x 45. 
Fig. 73. Antenna, with setae omitted. x 30. 


Nanostreptus orthacanthus, sp. nov. 


Fig. 74. Collum, ventrolateral view. x17. 


NI 
On 


Fig. Ends of collum and succeeding two tergites, ventral view. x 17. 


Fig. 76. Cardo of mandible, ectal view. x 45. 


Soutn AMERICAN DIPLOPODA Pate XI 


106 University of Michigan 


Pirate XII 


Nanostreptus orthacanthus, sp. nov. 


Fig. 77. Gonopods of male, anterior view. x 22. 
Fig. 78. Right gonopod, posterior view. x 22. 
Fig. 79. Leg of sixth segment of male. x 45. 
Fig. 80. Caudal end of body, dorsal view. x17. 


Fig. 81. Anal scale. x 17. 


Nanostreptus astix, sp. nov. 


Fig. 82. Collum, lateral view. x17. 
Fig. 83. Outer face of cardo of mandible, in outline. x 45. 


Fig. 84. Anal scale. x17. 


SoutH AMERICAN DIPLOPODA it PLATE XII 


108 University of Michigan 


Pirate XIII 


Nanostreptus astix, sp. nov. 


Fig. 85. Caudal end of body, dorsal view. x17. 


Nanostreptus gracilior, sp. nov. 
Fig. 86. Collum, etc., lateral view. x 30. 
Fig. 87. Anal scale. x60. 
Epistreptus eustriatus, sp. nov. 
Fig. 88. Collum, etc., lateral view. x13. 
Fig. 89. Caudal end of body, ventral view. x 13 
Spirostreptus atoporus, sp. nov. 


Fig. 90. Collum, lateral view. x 17. 


Fig. gt. Anal scale. x17. 


Soutn AMERICAN DIPLOPODA PLATE XIIB 


(o>. 


110 


Fig. 


. 97. 


University of Michigan 


Prate XIV 


Orthoporus etholax, sp. nov. 


Collum, ete., lateral view (male). x 13. 


~ Collum and adjacent parts of female, lateral view. 


Anal scale of female. x15. 


Left gonopod of male, anterior view. xX I?. 


Left gonopod, posterior view. X17. 


Orthoporus walkeri, sp. nov. 


Anal scale. x17. 


x FS 


ei 


Soutn AMERICAN DIPLOPODA PLATE XIV 


LIZ 


University of Michigan 


PLATE XV 


Orthoporus walkeri, sp. nov. 


l‘ig. 98. Collum, etc., of female. x 13. 


Tig. 


"ig. 


lig. 


Tig. 


Tig. 


Io]. 


102, 


103. 


Orthoporus gaigei, sp. nov. 


Collum of female. x 13. 
Collum of male. x 13. 


Right gonopod of male, viewed from a little laterad of 


directly in front. x17. 
Tip of right gonopod of a variant male, anterior view. x17. 


Tip of left gonopod of same variant male. x 17. 


SoutH AMERICAN DIPLOPODA PLATE XV 


114 


Fig. 


Fig. 


Fig. 
Fig. 


Fig. 


Fig. 


104. 


100. 


107. 


108. 


100. 


University of Michigan 


PLATE XVI 


Orthoporus gaigei, sp. nov. 


Anal scale. x 17. 


Orthoporus foliatus, sp. nov. 


Collum, etc., of female, lateral view. x 13. 


Rhinocricus amblus, sp. nov. 


Caudal end of body, lateral view, male. x17. 
The same, dorsal view. x17. 


Gonopods of male, anterior view, the left posterior gonopod 


omitted. x 30. 


Microspirobolus tridens, sp. nov. 


Gonopods of male, paratype, anterior view. x 45. 


SoutH AMERICAN DIpPLopopa 


PLATE XVI 


54s Barby University of Michigan 


Pirate XVII 


Microspirobolus tridens, sp. nov. 


Fig. 110.. Posterior gonopod, caudal view. x75. 


Pycnotropis colombiensis, sp. nov. 


Fig. 111. Keels of collum and succeeding two tergites. x 13. 
Fig. 112. Left gonopod, caudoventral view. x 30. 


Fig. 113. Right gonopod, ectal view. x 30. 


Pycnotropis cylindroides, sp. nov. 


Fig. 114. Tenth right keel of male (type). x13. 


Fig. 115. Seventeenth, eighteenth, and nineteenth right keels in out- 


line. x13. 


Fig. 116. Left gonopod, caudoventral view. x 30. 


Soutn AMERICAN DipLoPopa Pirate XVII 


118 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


117. 


118. 


119. 


120. 


E21. 


122. 


123. 


University of Michigan 


Pirate XVIII 


Rhyphodesmus amphelictus, sp. nov. 


Distal end of gonopods, ventral view. x17. 


Left gonopod, ectal view. x17. 


Aphelidesmus guianensis, sp. nov. 
Lateral portion of collum and second tergite, laterodorsal 
view. x13. 


Fourth and fifth keels, male, view a little lateral of dorsal. 


Be 1st 
Anal scale: x13: 


Right gonopod, ectal view. 


Trachelodesmus angulatus, sp. nov. 
Left ends of collum and second tergite, female, laterodorsal 
view. X 33. 
Seventeenth right keel. x 33. 


Left sternal spines of seventeenth and eighteenth segment:, 


ectal view. x45. 


Pirate XVIII 


SoutH AMERICAN DIPLOPODA 


120 University of Michigan 


PLATE XIX 


Trachelodesmus ancylophor, sp. nov. 


Fig. 126. Left ends of collum and second tergite, view a little lateral 


of dorsal. x 33. 


Left keel of fifteenth segment. x 33. 


yt 
dQ 

— 

to 

aT 


Dromodesmus longipes, sp. nov. 
Fig. 128. Left ends of collum and second tergite in outline (tubercles, 
etc., omitted), dorsal view. x 13. 
Fig. 129. Tenth left keel, in outline, dorsal view. x 13. 


Fig. 130. Right side of caudal end in outline, dorsal view (setae of 


cauda omitted). x 13. 
Fig. 131. Antenna (male type), in outline. x 13. 
Fig. 132. Left gonopod, ventral view. x 33. 


Fig. 133. Anal scale. 


SoutH AMERICAN DIPLOPODA PLATE XIX 


122 University of Michigan 


PLATE XX 


Dromodesmus longipes, sp. nov. 


‘Fig. 134. Leg of fourth to last pair of male type, in outline. x 13. 


Fig. 135. Left gonopod, ectal view. x 33. 


Colombodesmus catharus, sp. nov. 
Fig. 136. Left ends of collum and succeeding two tergites in outline, 
the tubercles, etc., not represented. x 17. 
Fig. 137. Tenth left keel in outline. x 17. 
Fig. 138. Left side of caudal end of body, dorsal view, in outline. x 17. 


Fig. 139. Right sternal spines of last two pediferous segments, with 


bases of legs. x 30. 


Fig. 140. Anal scale. x 30. 


SoutH AMERICAN DIPLOPODA PLATE XX 


124 


Tig. 


Fig. 
Fig. 
Fig. 


Fig. 


14]. 


Bee. 8 
. F4S, 


. 144. 


145. 
146. 
147. 


148. 


University of Michigan 


Pirate XXI 


Colombodesmus catharus, sp. nov. 


Penult leg in outline (setae and granules omitted). 


Colombodesmus lygrus, sp. nov. 


Tenth keel, dorsal view. x17. 
Penult leg (granules and setae omitted). x 30. 


Eighteenth and nineteenth left keels. x 17. 


Cormodesmus hirsutellus, sp. nov. 


Collum and second tergite, dorsal view. x 30. 
Antenna. x 30. 
Eleventh left keel. x 30. 


Anal scale. x 45. 


X 30. 


SoutTH AMERICAN DIPLOPODA PLATE XX} 


126 


Tig. 


Fig. 


Fig. 


Tig. 


Fig. 


Fig. 
Fig. 


Fig. 


140. 


University of Michigan 


Pirate XXII 


Cormodesmus hirsutellus, sp. nov. 


Gonopods of male, type, ventral view. x 30. 


Alassodesmus reductus, sp. nov. 


Thirteenth right keel. x 30. 


Sternal spines and bases of adjacent legs of last two pedif- 


erous segments, lateral view. xX 30. 
Anal scale. x75. 


Leg of one of last pairs, with setae omitted. x 45. 


Trichomorpha tuberculosa, sp. nov. 


Tenth keel. x17. 
Eighteenth and nineteenth keels. x 17. 


Seventh leg of male, with setae omitted. x17. 


SoutH AMERICAN DIPLOPODA | Peate- XXIT 


128 University of Michigan 


PLATE XXIII 


Trichomorpha tuberculosa, sp. nov. 


Fig. 157. Left gonopod, mesal view. x75. 


Fig. 158. Gonopod, ventroectal view. x 75. 


Trichomorpha rugosella, sp. nov. 


Fig. 159. Anal scale. x 45. 
Fig. 160. Left gonopod of male, ventral view. x 75. 
Fig. 161. Left gonopod of male, mesal view. x 75. 


Vig. 162. Tip of left gonopod, ectal view. x 75, 


Pirate XXIII 


SoutH AMERICAN DIPLoPoDpA 


158 


130 


Fig. 


Fig. 


Fig. 
Fig. 
Fig. 


Fig. 


e103: 


. 164. 


165. 


166. 


167. 
168. 
169. 


170. 


University of Michigan 


PLATE XXIV 


Trichomorpha rugosella, sp. nov. 


Seventh leg of male, with setae omitted. x 30. 


Eighteenth and nineteenth left keels of male, type. 


Trichomorpha setostor, sp. nov. 


Sixth keel. x 30. 


Eighteenth and nineteenth keels. x 30. 


Trichomorpha eutyla, sp. nov. 


Left end of tenth tergite. x 30. 
Seventeenth to nineteenth keels, in outline. x 30. 
Right gonopod, ventral view. x75. 


Right gonopod, ectal view. x75. 


x 30. 


SoutH AMERICAN DIPLOPODA PLATE XXIV 


Fig. 


Fig. 


Fig. 


Fig. 
Fig. 
Fig. 
Fig. 


lig. 


r71. 


173. 


University of Michigan 


PLatE XXV 


Trichomorpha eutyla, sp. nov. 


Seventh leg of male. x 30. 


Trichomorpha paurothrix, sp. nov. 


Right half of seventeenth tergite. x 30. 


Trichomorpha angulella, sp. nov. 


Right ends of Gehtocnth and nineteenth tergites. x 45. 
Trichomorpha eusema, sp. nov. 

Tenth keel, etc., of male type. x17. 

Eighteenth and nineteenth keels, etc., male type. x17. 

Seventh leg of male (type). x 30. 

I,eft gonopod, ventral view, type. x7:. 


Left gonopod, ectal view. x75 


SoutH AMERICAN DIPLOPODA Puate XXV 


171 


134 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


183. 


184. 


University of Michigan’ — 


PLatE XXVI 


Chondrodesmus tamocalanus, sp. nov. 


Twelfth right keel of type, male. x17. 
Eighteenth left keel of same. x17. 
Tenth keel of a male paratype. x13. 


Right gonopod, anteromesal view. x 45. 
Chondrodesmus cerasinopes, Sp. nov. 

Eleventh keel, in outline. x 13. 

Seventeenth to nineteenth left keels. x 13. 


Chondrodesmus virgatus, sp. nov. 


Tenth left keel. x17, 


Seventeenth left keel. x17. 


Prats XXVI 


SoutH AMERICAN DIPLoPoDA 


136 


Fig. 


187. 


. 188. 


. 180. 
. 190. 
. BGA, 
892. 


mrgs: 


. 104. 


University of Michigan 


Pirate XXVIII 


Chondrodesmus rugosior, sp. nov. 


Tenth keel. x13. 

Left gonopod, antedomesal view. x 30. 
Zigwadesmus guiananus, sp. Nov. 

Third, fourth and fifth keels. x17. 


Tenth keel. x17. 


Thirteenth and fourteenth keels, lateral view. 


Anal ‘scale. =x 17: 


Cauda, etc., lateral view. «17. 


Zigwadesmus modestus, sp. nov. 


Anal scale. x 30. 


X17. 


Soutn AMERICAN DIPLOPODA Puateé XXVII 


. iffy oo 
pe 


—— 


——<-. 


138 


Fig. 


Fig. 
Fig. 
Fig. 


Tig. 


Fig. 


Fig. 


195. 


196. 
197. 
108. 


199. 


201. 


(University of Michigan 


PLarE XXVIII 


Zigwadesmus guiananus, sp. nov. 


Last tergite, dorsal view. x17. 


Zigwadesmus modestus, Sp. nov. 


Fourth and fifth keels. x 30. 


Tenth keel. x 30. 


Thirteenth and fourteenth keels, lateral view. 


Last tergite, dorsal view, with setae omitted. 


Arionus ulophilus; sp. nov. 


Anal scale. x75. 


Tenth keel. 


X 30. 


X 30. 


Prats XXVIII 


SoutH AMERICAN DIPLOPODA 


140 University of Michigan 


Prateh XXIX 


Arionus ulophilus, sp. nov. 
Fig. 202. Head with collum and second tergite, ventroanterior view. 
X 30. 


Fig. 203. Collum and succeeding three tergites, lateral view, in out- 


line. x 30. 


Fig. 204. Caudal end of body in outline, dorsal view (tubercles of 


general surface omitted). x 30. 
Fig. 205. Gonopods, ventral view. x 45. 


Fig. 206. Gonopod, mesal view. x75. 


Guianonus ectoporus, sp. nov. 
Fig. 207. Last three tergites. x 45. 
Fig. 208. Anal scale. x75. 


Fig. 209. Gonopods of male, caudal view. x75. 


Soutn AMERICAN Drproropa PLAT XXX 


142 University of Michigan 


PLAGE OOo 


Guianonus ectoporus, sp. nov. 


Fig. 210. Right side of anterior end of body in outline, dorsal view. 


x 45. 


Fig. 211. Tenth and eleventh keels in outline. x 45. 


Agnurodesmus thrixophor, sp. nov. 


Fig. 212. Collum and the three following tergites, lateral view. x 17. 


Cliodesmus cryptopygus, Sp. nov. 


Fig. 213. Head, collum and three following tergites, lateral view. x 30. 
Fig. 214. Anal end of body, dorsal view. x 30. 
Fig. 215. Anal end of body, ventral view. x 30. 
Fig. 216. Gonopods in situ, ventral view. x75. 


Fig. 217. Right gonopod in situ, mesal view. x75. 


<outH AMERICAN DIPLOPODA PrATreE XXX 


juan te Khe AG 
REL WISis Oo, AERTS? SID 
, - - Wy - ie : el 


2 — 
* 2 4 


4 ? ° - / 
f 
7 
' 

‘ de, 4 

4 x 

s ; J 
f 
? 
- 5 P 
«* 
’ * 
. “ 
ae ’ ». 
- 
ca 2 : 
“> . 

“4 
34 
we 
4 ‘ 

rf . 's 

Pree he . ~ 


NUMDER 134 FEBRUARY I0, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arsor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NEW SPECIES OF ARCHAEOGOMPHUS 
(ODONATA) 
By E. B. WILLIAMSON 

In Occasional Papers of the Museum of Zoology, Univer- 
sity of Michigan, Number 59, June 24, 1918, and Number 63, 
January 5, 1919, I have described a new gomphine as Archaeo- 
gomphus hamatus. ‘This paper is supplementary to the two 
previously published papers. 

In Paper Number 63 it was pointed out that species referred 
to Agriogomphus by Needham, Ris and Williamson did not 
belong to that genus, and the new genus Archaeogomphus was 
proposed for wings figured by Needham, species not named; 
for Agriogomphus infans Ris, known from two females; for 
Agriogomphus hamatus Williamson, known from both sexes, 
and for an unnamed female from Colombia described by 
Williamson. 

The University of Michigan Expedition to Venezuela col- 
lected a single male and female of a species evidently the same 
as the unnamed Colombian female. The species is distinct 
and is therefore here described and named. 


2 University of Michigan 


Archaeogomphus furcatus, new species 


Abdomen, male 24, female 24; hind wing, male 18.5, female 
19-20. 

Male—Like A. hamatus except as noted below. Labrum 
with only a trace of a diffuse median brown spot; frons above 
slightly brownish, no defined triangular median spot; occiput 
rounded off posteriorly as in hamatus, and at either end a 
small, scarcely discernible knob where the horns of the female 
occur. 


Abdominal segment 2 slightly darker above than laterally, 
except at base, but no distinctly defined areas except the pos- 
terior black edge on the auricle; posterior suture black or 
nearly so (brown in hamatus) ; 3 as in hamatus, but with the 
apical brown not evident; likewise on 4-6 the apical dark is 
evident only as an ill-defined darkening on each segment, suc- 
cessively darker posteriorly and occupying about one-third of 
6; the dark ring in the basal pale area of each of segments 4-6 


is narrow as in 3, and much narrower than in hamatus. 


Second and third femora brown above, little if any darker 
at apex; tarsi brown, marked yellow as described for hamatus. 
(A re-examination of hamatus shows that in at least one case 
the tarsi are largely yellow, only the joints and the apex of 
the ‘last segment being brown or black.) 


Second hamule with the apical hook slightly more recurved 
than in hamatus and, the ventral border of the hamule, anterior 
to the hook, slightly elevated and rounded; not as straight as 
in hamatus. 

The above description is based on a single male of furcatus. 
The body colors of these gomphines are in general not in sharp 
contrast and postmortem changes operate still farther to oblit- 
erate the obscure pattern lines; so color differences between 


Occasional Papers of the Museum of Zoology 3 


hamatus and furcatus at present should not be relied on in 
separating the species. 


Female—In the teneral Bejuma female the body markings 
are all obscure and ill defined. The sutures between the abdom- 
inal segments are black; there is a narrow black ring at about 
two-fifths the length of 3, one-third the length of 4, and one- 
fourth the length of 5-7; the base and apex of each:of 3-7 
with more or less distinct traces of darker, especially the apex 
and laterally ; 8-10 dull, slightly darker than the segments basal 
to them. 

In the adult Cristalina female, which has been previously 
described (Paper Number 59, loc. cit.), in lateral view abdom- 
inal segments 3-6 have each an inferior dark stripe from base 
to and very slightly beyond the postbasal dark ring and the 
apical black on 3-5 is produced anteriorly along the ventral 
border more than half the distance from the apex to the post- 
basal dark ring; on 6 the black is as extensive dorsally as 
laterally. 

As in the case of the male, the description of body colors 


must be interpreted cautiously. 


Male and female—Wings similar to hamatus. Antenodals 
front wing, male 1o, female 11 (75%) or 10 (25%); hind 
wing, both sexes, 9. Postnodals front wing, male 5, female 5 
(75%) or 6 (25%); hind wing, male 5, female 5 (75%) or 6 
(25%). Inthe front wing the number of cells on the anterior 
side Cu, which do not reach M, is 1, and in the hind wing it is 
I in the male and in'the female 1 (25%) or 2 (75%). In 
the hind wings the number of cells posterior to Cu, and distal 
to the postanal cell which do not reach the posterior wing mar- 
gin is none or a single very small cell in the male; in the 


female there are 2 cells (the Bejuma female) or 3 (the Cris- 


4 University of Michigan 


talina female, misinterpreted as 4, due to crossvein shifting, 
in the original description, Paper Number 59, loc. cit.).* In 
the triangles of the front wing of the male and both females 
the distal part of the anterior side is equal to about two-fifths 
the combined lengths of the proximal and distal parts of the 
anterior side of the triangle; and likewise in the hind wings of 
all the specimens {the distal part of the anterior side equals 
about one-fourth the combined lengths of the two parts of 
the anterior side. 

The following brief color notes were made of the type male 
at the time of capture: Eyes brilliant bright green above, 
sharply pale brownish green below; in front with a large light 
yellowish brown pseudopupilla surrounded with green. Tho- 
rax light brownish yellow, marked with black and brown. 
Abdomen light brownish yellow and black; 7, apical to the 
median transverse carina, light yellowish green; 9-10, light 
reddish brown. 

Other descriptive notes on A. furcatus are included in the 
final part of this paper where the genus is described and the 
three known species are briefly discussed. 

Habitat: Colombia and Venezuela. Described from the 
type male, Bejuma, Carabobo, Venezuela, February 18, 1920, 
the allotype female, February 15, 1920, and a female, Crista- 
lina, Antioquia, Colombia, February 14, 1917, all in coll. E. 
B. W. 


On February 15, 1920, our collecting party went west out 


*In hamatus I find one male wing with none and one female wing 
with one only, thus differing from my original description (Paper 
Number 59, Joc. cit.), In the male and female wings figured (Plate 
II, Paper Number 59, loc. cit.) there is one cell in the male and two 
in the female wing. Caution is required in interpreting this character 
if cross-veins posterior to the subtriangle are switched about as is 
sometimes the case. 


Occasional Papers of the Museum of Zoology 5 


the main street of Bejuma to the Bejuma River and followed 
down stream all day. ‘The river is about fifteen to thirty feet 
wide, in a sand or gravelly bed, shallow pools alternating with 
gentle ripples, and the banks, immediately adjacent to the 
stream, are largely covered with wild cane. ‘There are less 
frequent growths of small trees and occasional open spots of 
grass or solid spreads of convolvulus. Where we left the river 
in the afternoon some round, grass-covered hills lay on the 
left bank, and between these hills and the river was a small, 
poorly tended and sickly coffee planting with a few large shade 
trees scattered through it. At the foot of the hills at the edge 
of the coffee planting were a few small swampy spots. 

After the day’s collecting we found in Jesse Williamson’s 
material a single slightly teneral female Archaeogomphus which 
he failed to recognize when it was captured, but which he 
knew he had collected somewhere on the river. On February 
18 I returned to the river to search for more specimens of 
the species. Cutting across country, I struck the river at the 
coffee planting, which, in view of our observations of A. 
hamatus in Colombia, seemed to me the most likely spot along 
the river for specimens of the genus. A thorough search 
over the entire coffee planting yielded nothing, so I started 
up river, working carefully adjacent cane patches, dry woods, 
a small banana field, and the broad expanses of waist-high 
convolvulus leaves, but without success. About two miles of 
the river was thus worked and I then returned, working down 
stream as carefully as I had worked up, and arriving without 
success about 4:30 p. m. at the coffee planting. Here I found 
many small libellulines resting on the tips of the dead twigs 
of the coffee trees (really bushes). These libellulines were 


busily inspected for half an hour in the hope of detecting an 


6 University of Michigan 


Archaeogomphus among them, and about 5 o’clock I gave up 
the search and started home. Passing from the coffee planting 
near the river, I looked back for the last time and saw, resting 
on a dead twig tip about three feet from the ground, a small 
dragonfly whose wings were horizontal. instead of slightly 
drooping in the almost invariable small libelluline position. I 
retraced my steps, and as I approached I saw the separated 
eyes of a gomphine, which, after a few literally breathless sec- 
onds, was safely fluttering in my cyanide bottle. This speci- 
men is the type of the species. I worked over the coffee plant- 
ing several‘times in the next hour till the setting sun ended 


the day’s collecting, but not another Archaeogomphus was seen. 


Tur GeNus ARCHAEOGOMPHUS AND ITS SPECIES 

In view of the characters shown by the new species 
described above, the genus Archaeogomphus may now be 
defined as follows: Small, short-legged gomphines with the 
third femora reaching backward slightly beyond the base of 
abdominal segment 2. Venation simple; triangles, supratri- 
angles and subtriangles free, crossveins between M,_, and M, 
not specialized, triangle four-sided, stigma without brace-vein, 
basal antenodal of second series wanting, one cubito-anal 
crossvein in addition to the inner side of the subtriangle, one 
row of postrigonal /cells in both front and hind wings, anal 
area of the front wing one cell wide, two postanal cells in the 
hind wing, cells posterior to Cu, and distal to the postanal 
cells in the hind wing, which do not reach the posterior wing 
margin, none to four in the male and one to six in the female, 
anal triangle wanting in the male, but posterior margin strongly 
angled. Male with the abdominal appendages reduced and 


functionless as grasping organs, the dorsum of segment 10 


Occasional Papers of the Museum of Zoology 7 


armed with two dorsal basal hooks and the dorsal apex of the 
segment produced posteriorly in a long tapering snout; female 
vulvar lamina with a broad, short base and two long tapering 
branches which reach nearly to or slightly beyond the apex of 
segment 9, branches apparently flexible as, in the same species, 
they are parallel and contiguous or the apices may be separated, 
the apex of each branch curved outward and away from the 
other branch. 

With the description of A. furcatus in this paper three spe- 
cies of Archaeogomphus are now known. The wings of a 
male from Brazil have'been figured but not named by Need- 
ham. It is not impossible that this male is A. infans Ris 
described from two females, the only known specimens, from 
Argentina. The other two species, iamatus and furcatus, are 
known from both sexes. 

The females of the three species may be separated by the 
following brief key: 

1. Rear of occiput armed with two posteriorly directed spines or 


horns; two or three cells posterior to Cu, in the hind wing which 
do not reach the posterior wing margin.................. furcatus 


1’. Rear of occiput not armed with posteriorly directed horns..... a 


2(1’). One to three cells posterior to Cuz in the hind wing which 
do not reach the posterior wing margin.........0....e.0.. hamatus 


2’. Six cells posterior to Cu, in the hind wing which do not reach the 
RE MRI PIER os ows « «ok w ac chee Boe eee wee c* infans 


It is certain that the venational character of infans employed 
above and based only on Ris’s figure of one wing will prove 
variable, but I believe the character, when the limits of varia- 
tion have been determined, will prove sufficient for the separa- 
tion of infans and hamatus. In the coalescence of the five 
veins at the posterior angle of the triangle in the hind wing 


of infans and the separation of these veins into two groups, 


8 University of Michigan 


three anterior and two posterior, in hamatus, we have another 


venational character probably of specific value. 


It is assumed in this paper that the Cristalina female pre- 
viously described (Paper Number 59, loc. cit.) is conspecific 
with the allotype of A. furcatus from Bejuma, as no charac- 
ters for separating these two females have been detected. How- 
ever, the Magdalena basin in which Cristalina lies and the 
Orinoco basin in which Bejuma lies are widely separated, and 
it is not impossible, though I believe it is improbable, that 
when males from the Magdalena basin are available, they may 
be found to differ from those of the Orinoco basin. In fact, 
with such scanty material no prediction as to the number of 


species occurring in each region is possible. 


In the allotype female of furcatus ‘the postoccipital horns 
are as figured for the Cristalina female (Paper Number 59, 
loc. cit., Plate I, figure 13), but due to the teneral condition 
of the allotype the horns are bent and slightly crumpled against 
the prothorax as the head is turned one-fourth around with its 
dorsal surface to the side. 


The vulvar lamina is likewise identical in the two and is 
very similar to that of hamatus (figure 11, loc. ctt.), except 
that in furcatus the branches are more slender, slightly longer, 
the outer edges more nearly parallel, and they come off from 
the base more abruptly, or, to express this last point differ- 
ently, each branch at the base is narrower in furcatus than in 
hamatus. ‘The vulvar lamina of infans has not been figured, 
but, as described, it is of the same general character as it is in 
hamatus and furcatus. 


The known males of ;Archaeogomphus are beautifully sep- 
arated by the outline of the first joint of the penis (seminal 


vesicle) in posterior view, as shown in figures 3 and 4 accom- 


Occasional Papers of the Museum of Zoology 9 


panying this paper. Other characters are found in the tenth 
segment (compare figures 3 and 4, Plate I, Paper Number 59, 
loc. cit., with figures I and 2, in this paper). In furcatus the 
dorsal hooks on abdominal segment I0 are more slender and 
are directed posteriorly, while in hamatus the hooks are directed 
interno-posteriorly. In dorsal view the snout-like apex of I0 


is broader in furcatus than in hamatus. 


NUMBER 135 FEBRUARY 25, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arbor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


THE MOLLUSCA COLLECTED BY THE UNIVER- 
SITY OF MICHIGAN-WALKER EXPEDITION IN 
SOUTHERN VERA CRUZ, MEXICO, IV 


By H. Burrincton BAKER 


The introduction and parts I, II, and III of the present 
paper appeared as number 106 of this series (Feb. 18, 1922). 
The habitat references (preceded by H) are explained in the 
introduction. Attention is called to the following errata in 
the preceding parts: 


Page 8, 5 lines from bottom: (?) to precede U. plicatulus. 


Pages 10, 11, 12: &. p. crocodilorum, not crocodilarum 
(corrected by L. S. Frierson, in letter). 

Pages 24, 25, 27: Arotonaias, not Artonaias (correction 
due to L. S. Frierson). 

Page 47, 6 lines from bottom: Drymeus, not Drymacus. 

Pages 48, 49: £. elegantulus, not elegantula. 


Page 56, 5 lines from bottom: T. hornii, not koriiit. 


2 University of Michigan 


SPHAERIIDA: 
Pisidiuim atlanticum Sterki. Sixteen specimens from near 
bank of La Laja, buried in wet humus and leaves at edge of 
forest pools (H, v, a). The growth-wrinkles on these speci- 


mens are considerably coarser than is usual in the species. 


PLANORBIDA; 


Planorbis cultratus d’Orbigny. One specimen; on roots of 
water plants, northern corner of Lake Catemaco (H, vii, d). 
More or less intermediate between this species and P. sumi- 
chrasti C. and F., this specimen appears closer to the former. 
It is somewhat higher than typical P.'cultratus, and the cari- 


nation of the outer lips of the aperture is not quite so acute. 


Planorbis isabel “Morelet” Sowerby (1879). 

P. ysabelensis C. and F. (1879). 

Eleven specimens; 10 from pools in ‘burnt-over area (H, 
v, b); 1 from rocks in shallow water near shore of Laguna de 
Catemaco (H, vii, d). A peculiar species, with each whorl 
extensively enveloping the preceding one. 

Planorbis retusus Morelet? A single small, broken speci- 
men, but the growth and spiral riblets are well marked; from 
lowland forest ponds along La Laja (H, v, a), perhaps drifted 
in from La Laja itself (H, vi, a). 

Planorbula orbicula (Morelet). ‘Two rather small speci- 
mens; from Laguna de Catemaco'(H, vii, d). 

Planorbula obstructa (Morelet). Five specimens; from 
lowland forest pools (H, v, a) and pools in burnt-over region 
(H, v, b). These correspond very well with Crosse and 
Fischer’s figures. 

Planorbula dentiens cannarum (Morelet). Nine specimens; 


from roots of water-plants, Laguna de Catemaco (H, vii, d). — 


Occasional Papers of the Museum of Zoology 3 


Although most of the specimens possess the typical, internal 
callus, none have developed the teeth. 


PHYSIDA 
Physa spiculata Morelet. Seven specimens; from pools in 
lowland forests (H, v, a) and in the burnt-over region (H, v, 
b), and from roots of water-plants (Pistia) along the shore of 
Laguna de Catemaco (H, vii, d). 


ANCYLIDA; 
Uncancylus sp.? Several specimens that appear to belong 
to this genus were found on rocks, etc., near shore of Laguna 
de Catemaco (H, vii, d). They have been sent to Dr. Bryant 


Walker for examination. 


PUPILLIDA 
Sterkia bakert Pilsbry (1921). One specimen, the type, 
from ground in lowland forests (H, i, a). 


Pupisoma dioscoricola insigne Pilsbry. ‘Twenty-three speci- 
mens; from leaves of trees in lowland forests (H, i, b) and 
the savannah brush (H, ii, b). (Identified by Dr. Pilsbry.) 


SUCCINEIDZ 
Succinea virgata von Martens. One adult and 4 young 
specimens ; on floating debris in Arroyo Hueyapam, near mouth 
of La Laja (H, vii, a). The adult measures: 
Altitude Greatest diameter Heightaperture Diameter aperture 
12.3 mm. 59 (7.3 mm.)1 67 (8.3 mm.)1 47 (5.8 mm.)1 
From the description and figures, I cannot separate this 
species from the more southern S. guatemalensis Morelet, 
1As in the previous parts of this paper, the first dimension is 
given in millimeters, while the remainder are each expressed, as a 


percentage, in terms of the first; these ratios are followed by the 
actual measurements in millimeters (in parentheses). 


4 University of Michigan 


which has the priority. Specimens from Guatemala also are 
in the A.W. <5, P-. 
Succinea virgata microspira C. and 'F. (1878). 


S. uv. pueblensis C. and F. (1878), not S. pueblensis C. and F. 
(1877, 1878). 


Eleven specimens; from the banks of La Laja (H, vii, a), 
and from wet place with water oozing out over rocks, on the 


shore of Laguna de Catemaco (H, vii, d). Examples measure: 


Greatest Height Diameter 
Altitude diameter aperture aperture 
Hae avai aoe cae tm 62 (7.7 mm.) 73 (9.0 mm.) 50 (6.2 mm.) 


ee yit cche = seasieS anim: 63 (7.2 mm.) 75 (8.6 mm.) 50 (5.7 mm.) 

The Latin description of Crosse and Fischer (1878) gives 
the name of this variety as var. (beta) microspira (p. 659), 
but the French description gives it as var. (beta) pueblensis; 
under the remarks it is again called var. (beta) muicrospira. 
The plate does not use the varietal name (xxvii-3). Von Mar- 
tens (1900) re-describes the form as microspira n. 

Succinea brevis Dunker? One specimen; from grass on 
the cleared portion of the Hacienda de Cuatotolapam (H, ii, c). 
This shell has quite the shape of a young S. virgata, but is 
solider and more opaque than are even the adults of that spe- 
cies. The growth-lines are heavy, so as to give the shell almost 
a ribbed appearance, while the spiral lines are also quite evident. 
The color (after stay in alcohol) is creamy and opaque, alter- 
nating in stripes with a rather dark amber. ‘The shell has 
almost 3 whorls, and there is a rather definite callus on the 


columellar margin of the aperture. 
OLEACINIDA 
Streptostyla irrigua similis Strebel. Five specimens; from 
the ground in the lowland forests (H, i, a). The following is 


the synonymy of the entire species, as it appears to me: 


Occasional Papers of the Museum of Zoology 5 


A. Streptostyla irrigua irrigua (Shuttleworth). Typical form. 
Spiraxis (Streptostyla) irrigua Shuttleworth (1852) ; 
Streptostyla 1. C. and F. (1870) and von Mart. (1891). 
Streptostyla cingulata Crosse and Fischer (1868, 1870). 


The typical forms are those with a stronger tendency toward 
impressed lines and costulae near the suture; cingulata represents the 
extreme of this phase. 


B. Streptostyla irrigua shutileworthi (Pfeiffer). 
Spiraxis shuttleworthi Pfr. (1856). 
Streptostyla sallei Crosse and Fischer (1868, 1870) ; Stre- 
bel (1878), etc. 

These are the larger shells which resemble, to a certain degree, 
Streptostyla lattrei (Pfr.). From the original description, I certainly 
agree with Strebel’s (1878, page 51) observation, ‘that true shuttle- 
qorthi of Pfeiffer seems closer to sallei and edwardsiana C. and F., 
and is not what is ordinarily known by that name. 


C. Streptostyla irrigua edwardsiana Crosse and Fischer (1868, 
1870), and others. 


This form is very close to the preceding, but is somewhat more 
atteiuate. 


D. Streptostyla trrigua ;similis Strebel. 
Streptostyla shuttleworthi C. and F. (1870), Strebel (1878, 
p. 18), von Martens (1891), etc. 
Streptostyla similis Strebel (1878). 


As used here, this includes the smaller thin-shelled forms, usually 
called typical shuttleworthi, and also the shells with less prominent 
spiral, but more prominent vertical, sculpture, which are Strebel’s 
typical similis. Strebel’s shuttleworthi and his similis are not dif- 
ferent geographical, or probably not even ecological, races, as he men- 
tions specimens of both, obtained in the same lot (p. 19). As the 
differential characters are very variable, I do not think that a new 
name is necessary for the form usually called shuttleworthi. 


E. Streptostyla irrigua ventricosa von Martens (1891). 
S. shuttleworthi ventricosa von Martens (1891). 
A slightly stouter form of what is here called similis. 
This whole group of forms, all of them described from 
around Cordova and Orizaba, may be nothing more than varia- 


tions or ecological forms of a northern subspecies of S. lattret 


6 University of Michigan 


(Pfeiffer) from Guatemala. The longer and heavier forms 
(true shuttleworthi) certainly approach quite closely that spe- 
cies.. However, in all of the specimens that I have seen, S. 
lattrei has the embryonic whorls smaller in proportion to the 
size of the shell, so that the apex appears sharper than in S. 
irrigua. Also, in the former, the suture is more deeply 
impressed, and jeach whorl appears as if shoved up over the 
preceding one, so that the embryonic whorls look as if they 
had broken loose along the sutural attachment and slumped 
down into the surrounding whorls. In addition, in the speci- 
mens before me (A. N. 8. P.), the columella is more nearly 
truncate in Jatirei than in irrigua, although the descriptions of 
differen’ authors disagree on this point. I have not seen speci- 
mens of Streptostyla quirozi Strebel (1878), but, from the 
description and figures, it appears to be more closely related 
to S. streptostyla (Pfr.) or cylindracea (Pfr.), which com- 
prise a divergent offshoot from the same lattrei stock. 

My specimens belong to the form similis, as used here. 
They all show some signs of spiral sculpture, but two have the 
vertical, impressed lines especially well-marked near the aper- 
ture, so as to approach typical similis of Strebel. Two others 
slightly approach typical irrigua, in that the vertical lines are 
accentuated ‘near the suture. All are rather small and thin- 
shelled, and have very faint, wavy varices of darker fulvous 
on the light amber, general color. The whorls appear to have 
a sutural border, but the lens shows this is simply due to the 
transparency of the shell. The largest has a trifle over 6 
whorls; it measures: 


Altitude Greatest diameter Height aperture Diameter aperture 
21.5 mm. 42 (gmm.) 77 (16.5 mm.) 19 (4mm.) 


Salasiella margaritacea (Pfeiffer) (1857). Four specimens 


(3 adult); from among leaves and humus on ground in low- 


Occasional Papers of the Museum of Zoology 7 


land forests (H, i, a). ‘They were associated with Streptostyla 
irrigua similis, Von Martens’ (1891) diagnoses this species 
as “‘lacvis,’ but his figure shows the plicatulations, and Pfeif- 
fer’s description (1859) distinctly calls attention to them. My 
specimens show quite definite growth-wrinkles, bounded 
toward the aperture by impressed lines, which extend up to 
within 2 whorls of the apex. Traces of very obscure, spiral 
striations are also visible on the body whorls. ‘The apical 
whorls are practically smooth, even under considerable mag- 
nification. ‘The color of the shells is whitish-horn. They are 


rather small; the largest have 5 whorls and measure: 


Altitude Greatest diameter Height aperture 
8.5 mm. 42 (3.8 mm.) 71 (6.0 mm.) 
8.2 mm. 44 (3.6 mm. ) 71 (5.8 mm.) 


This species has the shortest spire in the genus, and also 
has the least-shouldered whorls. Nearest it, in general shape, 
are S. guatemalensis and S. pulchella, which differ from it by 
their somewhat higher spires, more marked sutures, and the 
flatter, vertical sides of their apical whorls. 

S. brownt Pilsbry from Panama, S. guatemalensis Pilsbry 
from Guatemala, S. pulchella (Pfeiffer) from Chiapas, S. joa- 
quine Strebel from Vera Cruz, and S. hinkleyi Pilsbry from 
San Luis Potosi, form a series of apparently quite closely 
related species. As a group, they decrease in size quite regu- 
larly from south to north (lengths 12, 9.9, 10.5, 8.5, 8.5 mm. 
in the order named), while the spire increases in comparative 
prominence both north and south of the area near the southern 
boundary of Mexico (length of aperture divided by length of 
shell equals 60, 66, 67, 60, 50 per cent, respectively, in the order 
named). The proportion: between the greatest diameter and 


the altitude is quite the same (35 to 37 per cent) in all of 


8 University of Michigan 


these species, with the exception of S. guatemalensis, which is 
slightly broader (40 per cent). 

Euglandina decussata (Deshayes), near subspecies tenella 
(Strebel). ‘Ten specimens, nearly adult, and 21 juvenile speci- 
mens; from the ground and the leaves of the trees in the low- 
land forests (H, i, a and b); dead shells from the burnt-over 
area (H, ii, a); from the leaves of trees in the savannah brush 
(H, iii, b), and one dead specimen quite far from the nearest 
trees on the savannah grassland (H, iv); also from near the 
Laguna de Catemaco. This species is a rapidly moving form, 
which appears to cover most of the ground habitats, and goes 
quite high up into the trees. The juvenile specimens are espe- 
cially common on the leaves of trees. 


ACHATINIDA® 

Opeas beckianum (Pfeiffer) (1846). Seventy adults and 
some juveniles; mainly from the ground in the lowland jungle 
(H, i, a), but also from the ground in the savannah brush (H, 
iii, a) and from near Lake Catemaco. These are quite char- 
acteristic of the slender Vera Cruz form (cf. Pilsbry, 1906), 
but have very distinct costulations on the middle whorls and 
sometimes quite to the aperture. The apex, under magnifica- 
tion, shows minute, spiral lines in both this species and in O. 
gracile. A figure (Figure 6) of the radula is given; the mars 
ginals differ from the laterals, mainly in their reduced size. 

Leptinaria martensi (Pfeiffer) (1857). One adult and 8 
younger specimens that are probably this form; from the 
ground in the lowland jungles (H, i, a); also 1 young speci- 
men, from near Lake Catemaco, that is either this form or L. 
mexicanum (Pfr.). The columellar fold appears rather 
variable in these shells, and the vertical costulae are quite well 
marked. 


Occasional Papers of the Museum of Zoology 9 


BULIMULIDZ 
Bulimulus coriaceus (Pfeiffer) (1857). Two quite typical 
specimens, except for their grayish color, on the ground in a 
banana plantation at the edge of the jungle, near Lake Cate- 


maco. ‘The larger measures: 


Altitude Greatest diameter Heightaperture Diameter aperture 
19.8 mm. 52 (10.2 mm.) 49 (9.8 mm.) 34 (6.7 mm.) 


Bulimulus coriaceus, var. a. Seven adults and numerous 
younger specimens, from the Hacienda de Cuatotolapam, show 
the dark chestnut band poorly, or even lack it entirely. A few 
young shells of this form were obtained on the ground in the 
lowland forests (H, 1, a), but the majority were collected from 
the cleared land (H, ii, b) along the railroad track, in sugar- 
cane and corn fields, and even in the middle of a road, where 
they were observed, coming up out of the ground in consider- 
able numbers, during a rain-storm. ‘This is plainly a deep- 
burrowing form, that appears to occur in colonies, and which 
thrives under conditions of cultivation. Most of the specimens 
were quite typical in shape, but others were considerably more 


slender. Examples measure: 


Altitude Greatest diameter Heightaperture Diameter aperture 
15.8 mm. 56 (8.8 mm.) 48 (7.6 mm.) 36 (5.7 mm.) 
15.8 mm. 51 (8.0 mm.) 47 (7.2 mm.) 30 (4.8 mm.) 


The radula of this form is shown in figure 3. The entocone 
is represented by a distinct lamella on most of the inner teeth. 
The radular formula may be given as: 

I ESO 2 I 
C—;L—+—4—+4 —= 24— 1 — 24. 
3 2 3 4-3 
There is no marked differentiation of marginals, but the teeth 
toward the outside of the radula become smaller, tend to turn 


inward, and lose the entoconal lamella. 


10 : Universily of Michigan 


Bulimulus coriaceus, var. b. One shell from the grass on 
the savannah (H, iv) is very large and slender. It is buff in 
color and lacks the sutural band. If obtained in larger num- 
bers, it would seem to be worthy of at least subspecific recog- 
nition. It measures: 


Altitude Greatest diameter Height aperture Diameter aperture 
21.7 mm. 47 (10.1 mm.) 41 (8.8 mm.) 30 (6.4 mm.) 


Drymeus dominicus (Reeve) (1850). Nine adults and 5 
smaller specimens; from leaves of trees in the lowland jungles 
(H, i, b), in savannah forests (H, iti, b) and dead shells from 
ground in latter (H, iii, a) ; also one immature specimen from 
near Lake Catemaco, and numerous juveniles along Arroyo 
Hueyapam (H, ii, a). These shells are very variable in col- 
oration; one young specimen has all of the 5 bands present, 
but the upper 2 are broken into dots; another has the upper- 
most and lowest absent; two shells have bands 2, 3, 4 and 5, 
with 2 and 3 broken; while the remainder have only 3, 4 and 5, 
with 3 and sometimes 4 broken. An example measures: Alti- 
tude, 19 mm.; greater diameter, 55 (10.5 mm.). 

The jaw and radula were obtained from two dried speci- 
mens. ‘The radular formula (Figure 2) is 121—1—121. In 
one radula the rhachidian tooth bears two cusps and is quite 
symmetrical, but in the other its symmetry is usually disturbed 
by the presence of a third, smaller cusp. In all cases, the basal 
portion is long and slender. The remainder of the teeth can- 
not be sharply divided into laterals and marginals. Most of 
them have 4 cusps, but some of the inner three may have only 
3; teeth of both types occur in a single longitudinal row. In 
the outer portion of the radula the second cusp from the inside 
is larger and more spatulate than the others. The base of all 
of these lateral teeth is trapezoidal, and the posterior edge is 
almost as broad as the anterior. ‘The cusp-bearing portion 


Occasional Papers of the Museum of Zoology II 


has a very noticeable thickened prolongation of the outer ante- 
rior end. The outermost 2 or 3 teeth are somewhat reduced 
and have a larger number of cusps. On either side of the 
center the transverse rows extend obliquely forward, but curve 


so as to be almost transverse at their outer ends. 


Drymeus albostriatus (Strebel) (1882). Twenty-seven 
larger specimens and numerous immature ones; from the 
leaves of trees and brush (H, i, b; H, ii, a; H, iii, b), and also 
(dead shells) from the ground (H, i, a; H, ili, a). These 
shells are very similar \in form to the preceding species and 
occur with it. The coloration is always distinct. Only two 
specimens show, around the third whorl, 3 broad, spiral bands, 
broken by lines parallel to the growth-lines. Some of the 
shells are quite without brown markings, and even without 
distinct, milky-white varices, but the majority have quite dis- 
tinct, and rather numerous, chestnut-brown varices. In addi- 
tion, my specimens are smaller and somewhat heavier than 
those of D. dominicus; and the spiral striations are not as dis- 
tinct, above the greatest ventricosity of each whorl, as in that 
species. The columellar reflection also appears more extensive, 
and the umbilicus is simply a narrow slit. An example meas- 
ures: altitude, 14.5 mm.; greater diameter, 52 (7.5 mm.). 

The jaws and radulae were examined in two specimens. 
The radular formula (Figure 1) is 135—1—135. The rha- 
chidian tooth is broader and more recurved than in the pre- 
ceding species; the distal end bears 3 cusps, the middle one 
of which is larger than the others. The tooth is usually 
twisted to one side, so as not to be exactly symmetrical. Most 
of the lateral teeth bear 5 cusps, but some in the inner three 
rows have only 4, while some of the outer have 6; the varia- 


tion in the inner teeth may occur in a single longitudinal row, 


12 University of Michigan 


but the outer rows are usually more constant throughout their 
lengths. The body of each of these teeth is attenuate anteriad, 
so that the base is narrow. The cusps of each tooth are more 
nearly equal in size than in D. dominicus. Each half of the 
transverse rows is quite straight and extends obliquely anteriad 
from the center. 

For comparison, the radulae from two specimens of Dry- 
meus multilineatust were examined. ‘The rhachidian tooth is 
stout and short; the symmetrical tip is markedly recurved and 
bears three cusps. Most of the lateral teeth bear 4 cusps, but 
a reduction to 3 very commonly occurs in some of the inner 
three longitudinal rows. The enlargement of the second cusp 
from the inside, on each tooth, is even more noticeable than in 
D. dominicus. The body is attenuate anteriad somewhat as in 
D. albostriatus. ‘The transverse rows are V-shaped; each half 
extends obliquely anteriad from the center for a short distance 
and then curves slightly outward. The radular formula is 
144—I—144. 

Oxystyla princeps (Sowerby). ‘Twenty-two adult or nearly 
adult and 1 juvenile specimens; mainly from trees in the 
thick jungles (H, i, b), but also from the partially cleared 
places along Arroyo Hueyapam (H, ii, a) and the savannah 
brush (H, iii, b). Dead shells also picked up from the ground 
in the jungle (H, i, a) and the burnt-over places (H, ii, b). 
This species appears to be purely arboreal, and I have found it 
aestivating in cavities in the trees. 

These specimens are very variable in color and pattern, but 
no relations between these characters and the habitat could be 


made out. The variation may be analyzed as follows: 


1A.N. S, P. No. 88,779; Sugar Loaf Key, Fla.; J. B. Clark, May 
23, Ig2I. 


Occasional Papers of the Museum of Zoology 13 


A. Apex coloration: from dark chestnut to practically 


colorless. 


B. Varices. These dark bands, which appear to corre- 
spond to former resting periods in peristomal growth, are not 
very prominent in my shells, but as many as 3 occur on some 
of the specimens. A break in the color-pattern quite commonly 
occurs at the varices. For instance, in one shell the coloration 
of the major portion of the spire approaches trifasciata Pilsbry, 
although with less prominent color-pattern, but this changes 
abruptly, on the last whorl, to a pattern resembling crossei von 


Martens. 


C. General coloration. Almost white to light brown; usu- 


ally this background color becomes darker on the later growth. 


D. Flammulations. These may be almost black and quite 
sharply marked, or each may be surrounded by a diffusely col- 


” 


ored border, as if the colors had “run.” In some shells the 


coloration is so diffuse that the flammulations are quite indis- 
tinct. One shell is so diffusely colored that the axial bands 
can only be seen near the varices, while the last whorl shows 


four indistinct broad spiral zones of light brown. 


FE. Angle-spots. ‘These may not be identifiable; they may 
appear simply as three projections of each flammulation on 
the side toward the aperture; they may tend to become darker 
than the remainder of the color-pattern, so as to give the 
appearance of three broken spiral bands (trifracta Pilsbry) ; 
or they may join up completely (feriwssaci von Martens). 
When the flammulations are very indistinct or practically 
absent, but three well-defined spiral bands are present, the col- 
oration is like tricincta von Martens. None of my specimens 


belong to this last category, but one shell is even more diver- 


14 University of Michigan 


gent, as it has only the two lower spiral bands on an almost 
white background. Young shells (2 to 3 whorls) usually have 
flammulations above the angle and a spiral band below, but two 
of mine lack the flammulations and I believe correspond to 
the tricincta-like form. 

E. Shell-shape. One shell has a distinct scalariform ten- 
dency. This is the one mentioned above, with only two spiral 
bands of color, and the upper of these, which is usually hidden 
on all but the last whorl, is visible on all of the whorls, and is 
3 mm. above the suture on the penultimate one. This shell 
(last series of measurements) is more elongate than most speci- 
mens of O. longa (Pfeiffer). 


Three of the larger specimens measure: 


Altitude Greatest diameter Heightaperture Diameter aperture 
62.0 mm, 56 (35 mm.) 54 (33.5 mm.) 34 (21 mm.) 
55-5 mm. 59 (33 mm.) 55 (30.5 mm.) 36 (20 mm.) 
50.5 mm. 51 (26mm.) 50 (25.5 mm.) 32 (16 mm.) 


In two of the radulae examined from adult specimens of 
this species the peculiar aculeate tendency of some of the inner 
teeth is apparent (compare Pilsbry, 1902). In the youngest 
teeth of both specimens the central tooth is of this type. In 
one, the second lateral on the left side, and the inner three on 
the right are of this type; while in the other the inner three on 
the left side and the inner two on the right show this modifica- 
tion. On the other hand, all of the older teeth, towards the 
anterior edge of the radula, very closely approximate the nor- 
mal, rounded form (compare Pilsbry, 1902, for Liguus). In 
the very large radulae of O. princeps this can be seen to be due 
to wear (Figure 4), and the broken edges of the teeth are 
quite apparent in the middle portion of the radula. The outer 
teeth tend to lose the point more rapidly than do the inner, 
and the rhachidian tooth is the last to be worn down to the 


Occasional Papers of the Museum of Zoology 15. 


normal form, although sometimes evident breaks give excep- 
tions to this. 

Pilsbry (1895), in his classic on the Helicidz, has pointed 
out that modification of the radula tends to take place from the 
center out. However, in some groups of snails the radula 
tends to be concave, as is also the odontophore. ‘This is espe- 
cially noticeable in the Helicinide and Neritide, where there 
are two radular cartilages with a slit between. In these it 
would seem that the greatest stress would tend to come at two 
points some distance out from the center on either side. 

Coincident with this stress-tendency, the greatest adaptive 
modification in some groups seems to be, not at the very center 
of the radula, but at some distance out on either side. ‘Thus, 
in the Zonitidz the inner marginals appear to show the greatest 
modification along certain lines, while in the Helicinidz the 
lateral complexes are certainly the most highly specialized teeth. 
In those radula of this type which show progressive modifica- 
tion within the transverse row, as, for example, in EKuconulus 
and Guppya (Habroconus), the adaptive (7) specialization 
seems to become less toward the inside as well as toward the 
outside (compare Part III of present paper). 

In Oxystyla princeps the radula is concave at the center 
and wear appears to be greatest some distance out from the 
center on each side. On account of the high degree of modi- 
fication of all of the teeth, it is difficult to establish the posi- 
tion of greatest adaptive (?) modification, but it perhaps may 
be assumed to be near the position of greatest stress. For 
this reason, I am inclined to regard these lanceolate teeth as 
vestigial or atavistic rather than as nascent modifications. 

The variability in the numbers of these peculiar teeth and 


their occurrence in some individuals in most of the genera of 


16 University of Michigan 


Orthalicine appear to substantiate this idea. Vestigial or rever- 
sional characters are, as a general rule, more variable than new 
ones. In addition, the sporadic occurrence of these teeth in 
several genera appears to be most easily explainable by the 
hypothesis that they may represent an approximation to the 
ancestral laterals of this subfamily. In younger radulae (from 
specimens of three whorls) some of the outer teeth are more 
pointed than is usual in the older specimens, but the aculeate, 


lateral teeth do not appear to be much more common. 


VAGINULIDZE 

Vaginula moreleti Crosse and Fischer (1872). Occurs 
practically everywhere, but noted in greatest abundance in the 
grass along the edges of cleared (H, i, c) and partially cleared 
(H, ii, a) fields and in the sugar-cane plantings. It is appar- 
ently almost noctural in habits, as it was usually collected on 
wet mornings; but it was also found moving about on rainy 
days. In addition to these artificial habitats, it was collected 
from the lowland jungles, mainly on the ground (H, i, a), but 
also up on the vegetation (H, i, b); from the grass on the 
open savannahs (H, iv); and from the ground (H, ii, a) and 


low vegetation (H, iii, b) in the savannah brush. 


18 University of Michigan 


PLATE! 


The magnification of each of the figures is indicated by the hair- 


line under it; this represents an actual length of 50 microns (.05 mm.). 


Figure 1. Drymeus albostriatus. The central and the tips of the 
first laterals. The 1st, 3rd, 28th, 100th and 135th (outermost) teeth 
on the right side. The V-line shows the arrangement of a transverse 


row. 


Figure 2. Drymeus dominicus. Same arrangement as Figure I- 
The tip of a tricuspid central (from another radula) is shown between 


the central and the first lateral. 


Figure 3. Bulimulus coriaceus. Central and Ist, 3rd, 7th, 14th 
and 21st teeth on right side. 

Figure 4. Oxystyla princeps. Aculeate type of laterals from a 
single longitudinal row, to show the effects of wear. The 2nd left 
lateral of the 2nd, 7th, 14th, 35th, 49th, 56th and 77th transverse rows, 


counting from the anterior end of the radula. 


Figure 5. Drymeus multilineatus. Same arrangement as Figure 1. 
2] 4 > 5S 


Figure 6. Opeas beckianum. The central, the Ist, 7th, 14th, 20th 


and 21st (outermost) teeth of the right side. 


POODUAQ 


NUMBER 136 FEBRUARY 10, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arsor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


DESCRIPTION OF A NEW SPECIES OF PIPA 
FROM VENEZUELA 


By ALEXANDER G. RUTHVEN AND HELEN T. GAIGE 


A collection of reptiles and amphibians made in Venezuela 
by M. A. Carriker, Jr., for the Museum of Zoology, in the 
spring of 1922, contains several specimens of an apparently 


undescribed species of Pipa. 


Pipa parva, new species 


Diagnosis: The new species is easily distinguished from 
Pipa pipa (Lann.) by the narrower and less depressed head, the 
rounded snout and shorter fore leg. Thus, a P. pipa 32 
mm. in total length has a fore leg 19 mm. long, and the head 
at the angle of the mouth is 13 mm. wide, while a Pipa parva 
(adult) of the same length has a fore leg 12 mm. long, and 
the head is 8 mm. wide at the angle of the mouth. It differs 


both from pipa, and, as nearly as one can make out from 


2 University of Michigan 


the somewhat meager description, from snethlegee@, in the 
smaller size (adult pipa 165 mm., snethlegee 75 mm. 
parva 39 mm.), and in the absence of metatarsal tubercles and 
of tentacles and flaps on the head; and from snethlegee in 


having a longer hind leg. 


Type: Museum of Zoology No. 57,443; taken in a covered 
drain in the village of Sabana de Mendoza, Venezuela, by M. 
A, Carriker, jr.; May 5,. 1922. 


Description of Type: Head narrow; no trace of dermal 
flaps or tentacles on mouth and eyelids; tip of snout rounded, 
projecting. Fingers slender, subequal, ending in four appen- 
dages. Toes pointed, broadly webbed, no subarticular or meta- 
tarsal tubercles. The hind leg being carried forward along the 
body, the tarso-metatarsal articulation reaches half way between 
eye and tip of snout. Skin evenly tubercular above, more 
smooth beneath. 

Color (in alcohol) above blackish gray, obscurely spotted 
with darker. Grayish white beneath, with a few scattered dark 
spots on sides and throat. 

Total length, 39 mm.; width of head at angle of mouth, 9 
mm.; length of fore leg, 12 mm. 


Remarks: The five other specimens in the series were col- 
lected in the same village as the type. There is little variation 
in color and comparative measurements. They range from 30 
to 39 mm. in length, the width of the head is contained in the 
total length from 4 to 4% times (2!% to 3 times in adult speci- 
mens of pipa, and_ the tarso-metatarsal articulation, 
when the hind limb is extended along the side, reaches to the 
eye or just beyond it. One of the spectmens (39 mm. long) 


is a female with eggs embedded in the skin of the back. 


NUMBER 137 JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arbor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


THE MOLLUSCA COLLECTED BY THE UNIVER- 
SITY OF MICHIGAN-WILLIAMSON EXPEDITION 
IN VENEZUELA 


3y H. Burrinctron BAKER 
Parr I. Curacao 


' Curacac is a small island lying about 50 miles off the coast 
of Venezuela, in approximately 69 degrees west longitude by 
12 degrees north latitude. It has always been of considera- 
ble interest to conchologists, as its molluscan fauna, as far as 
known, appears to indicate West Indian, rather than South 


American affinities. 


As viewed from the steamer, it is a rocky, quite arid island, 
with a low cliff along the shore, at least at the northern eid. 
Back from the ocean are seen numerous cliffs, broken ridges, 
and occasional peaks, which reach an altitude of perhaps 30a 
to 400 meters. The north end of the island appears to %ave 
considerable brush, covering especially the higher Ailfs, but 


the vegetation is predominantly chapparal-like, with frequent 


y 


2 University of Michigan 


organ-pipe cacti. ‘The south-east end, near Willemstad, 1s 
lower, less broken, and noticeably more arid. From its color, 
much of the rock outcrop at the northern end appears to be 
metamorphic or igneous, but the south shore is entirely of a 
coral-reef formation. 

On March 28, and again on May 6, 1920, a few hours col- 
lecting at the east edge of Willemstad netted a considerable 
number of shells. The abundance of Cerion was astounding ; 
practically every bit of vegetation was almost covered at the 
base with estivating individuals. I estimated that there were 
about 50 specimens per square meter examined. 

The place collected is Schaarlo, just behind the lagoon 
east of the harbor. This hill is entirely composed of porous 
limestone, somewhat like that around Nassau in the Bahamas. 
The vegetation is very scanty and consists mainly of Opuntia- 
like cacti and prickly pears, with some of the larger organ-pipe 
cacti, mimosa and evergreen thorn-bushes reaching a height 
of about 6 meters. Extreme dessication is the predommant 
feature of the landscape, and I am told that it rains very 
rarely. All of the living shells collected were estivating 
either on the lower stems of bushes and cacti (mainly Cerion), 


or in crevices between and under the rocks and rock-frag- 


ments. 
ANNULARIID/® 


Tudora megacheilos (Potiez and Michaud). 


Quite abundant; mainly under bits of coral rock and in 
crevices, but rarely found estivating on the vegetation. This 
species is very variable, in color, size and sculpture. The 
males are considerably smaller than the females, and tend to 


be more brilliantly colored. The coloration may be classified 
as follows: 


Occasional Papers of the Museum of Zoology 3 


a. Ground-color: white to buff and old rose. 


b. Inside of aperture: buff to light chocolate-brown and 
scarlet, corresponding to the three extremes of ground-color. 
The stripes of the exterior also show through the shell. 


c. Tip of spire (embryonic whorls always absent in 
adults): purplish-black through dark chocolate-brown to the 
same shades as the ground-color. The shells with the pink- 


ish general coloration often have salmon or scarlet tips. 
d. Spiral bands: 


1. Number: the maximum is 5; one near the suture 
(only in one specimen); a group of 3 near the 
greatest ventricosity; and 2 around the umbilicus 
in the region of the parietal wall of the aperture. 
Commonly, only one band is present; this may be 
the central one of the peripheral group, or, even 


more commonly, the upper of the umbilical pair. 


2. Color: any shade of the series given for the tips, 
except the purplish-black. 


3. Continuity: either entire or broken into square dots; 
narrow and sharply marked, or wide and diffuse. 
Sometimes the central group of 3 fuse into a sin- 
gle broad band. 


e. Varices: the commonest form of banding consists sim- 
ply of diffuse axial varices, which vary in color as do the 
stripes. One specimen has yery distinct chocolate-brown vari- 
ces, with flammulations corresponding to the position of ihe 
spiral ridges; near the aperture band 4 is entire and.very prom- 
inent. 

Although variable in shape, all of the specimens are quite 


typical of T. meyacheilos. No specimens approaching 7. cos- 


4 University of Michigan 


tata (“Menke” Pfr.) (1846) are present in this lot, although 
specimens in the A. N. S. P., from both Curacao and Buen 
Ayre (Bland), approach the description of the latter. It will 
probably be found to be a local species or subspecies. 

The shape of the aperture is very variable. A tendency to 
form a broad, angular columellar reflection, and an almost can- 
aliculate, free upper angle is quite common. 

The apical whorls of young specimens are irregularly and 
lightly pitted. The remainder of the whorls have fine, close- 
set axial riblets, which are always quite regular. These are 
usually crossed by heavy spiral angulations; as many as 17 
may be present, or they may be so faint as to be practically 
absent. It is quite impossible to use this character to deter- 
mine the sectional position of this lot. One shell has only one 
spiral thickening, which runs around the greatest ventricosity 
and makes all of the whorls quite markedly angular. In addi- 


tion, a few shells are malleate. 


Representative examples measure: 


Greatest Height Diameter 

Altitude diameter aperture aperture 
male: 128mm. 70 ( 9.0mm.) 52 (67mm.) 45 (5.8mm.) 
female: i9. 8mm. 64 (12.7mm.) 50 (10.0mm.) 45 (8.9mm.) 


Chondropoma (?) raveni (Crosse) (1872). 

Ten dead specimens from rock debris. In some specimens 
the aperture is scarcely solute, while in others the entire last 
whorl is free. None of my specimens have the operculum 
(compare Henderson and Bartsch, 1920), but the shell char- 
acters are certainly closest to the group of Annularia lachneri. 
The apical whorls (figs. 1-6) are practically smooth, but show 
irregular anastamosing wrinkles under high magnification ; 
they are always eroded from adult specimens, but the total 


number of whorls must be about 9. 


Occasional Papers of the Museum of Zoology 5 


PUPILLIDA 
Pupoides simoni (Jousseaume) (1889) ? 


One fresh specimen. The peristomal callus is not well de- 
veloped ; the parietal callus is especially weak, in fact, prac- 
tically lacking. No angular tubercle nor reddish band shows 
on the throat of the aperture. The axial striations are distinct 
on the lower whorls. With 5% whorls, the specimen meas- 
ures: altitude, 3.9 mm.; greatest diameter, 46 (1.8 mm.). It 
appears closest to P. modicus (Gould) (1848), but may pos- 
sibly be a youngish specimen of P. marginatus nitidulus (Pfr.) 


(1839). 
ACHATINID<AE 


Leptinaria (Neosubulina) gloynii (Gibbons). Figs. I-1, 2, 3. 


Cionella gloynit Gibbons (1879), (gloynei auct.), Curacao. 
Neosubulina harterti Smith (1898), Buen Ayre. 
L. gloynii minuscula Pilsbry (1907). 


Thirty-seven specimens, from crevices of rock and among 
rock-fragments. From the variation in the lot before me, I 
can see no reason for the specific separation of any of the 
forms included in the above synonomy. From the figure, 
L. harterti appears less attenuate towards the apex. _E. A. 
Smith, from his list of species of Curacao, was apparently not 
acquainted with even the description of L. gloynit. 

The embryonic whorls of these specimens have close and fine 
axial striations, which become somewhat coarser on the later 
whorls. The egg is of quite large size, and the slender em- 
bryonic shell contains 114 whorls. The spiral lamellze on the 
columella extend back 4 whorls from the aperture, and the 
parietal one is present in young shells. My specimens show 
considerable variation in size, apparently not entirely due to 


age. An individual measures: 


6 University of Michigan 


Greatest Height Diameter 
Altitude diameter aperture aperture 
934 whorls 11.8 mm. 23 (2.7 mm.) 23 (27 mm;) I3Ges inane 
BULIMULIDA& 


Drymeus multilineatus (Say). Two dead, bleached specimens. 


UROCOPTIDz 
Microceramus bonairensis curacoana, new subspecies 
Figs. 1-4, 5 

Fighty-two specimens, some living, under bits of coral rock 
on outskirts of Willemstad, Curacao. 

Shell distinctly rimate, turrite to lanceolate; last 4 whorls 
almost equal in diameter, but the penultimate slightly the broad- 
est; earlier whorls gradually, or quite rapidly increasing in 
size. Color: light corneous to dark brown, usually marked 
with opaque, milky-white, irregular varices, which may be so 
extensive as to obliterate the corneous ground-color on the last 
whorls. Whorls: 9 to II, somewhat convex; the first half- 
whorl smooth and polished; the next ones with minute, regu- 
lar, closely spaced axial riblets, which are also quite well- 
marked on the corneous portions of the last whorls, but are 
almost obliterated by the thickening of the interstitial spaces 
on the opaque, whitish portions; these thickenings on the last 
whorls may develop several heavy, irregular, raised, spiral 
lines; last whorl with a quite well-marked, basal angulation, 
which gives the shell a peculiar subtruncate appearance when 
viewed from behind, and which is much more prominent in 
young shells. Aperture: nearly circular; peristome whitish, 
but slightly thickened and reflexed, incomplete on the parietal 
margin, most reflected on the columellar. Columella inside of 


whorls; cylindrical and quite slender, showing only a very 


vague spiral undulation. 


Occasional Papers of the Museum of Zoology 7 


I have not seen specimens of typical bonairensis (Ti. A. 
Smith, 1898), but it is certainly a smaller shell. The indi- 
viduals of curacoana figured show the extremes in variation 
of size and shape. They measure: 


Greatest Height Diameter 
Whorls Altitude diameter aperture aperture 
II 9.3 mm. 35 (3.2mm.) 27 (2.5mm.) 26 (2.4mm.) 
II 8.9 mm. 31 (2.7mm.) 22 (2.0mm.) 22 (1.9mm.) 
9 7.1mm. 41 (2.9mm.) 24 (1.7mm.) 25 (1.7 mm.) 
9 6.7 mm. 41 (2.7mm.) 25 (1.7mm.) 27 (1.8mm.) 
bonairensis 
8% 6.0 mm. 42 (2.5mm.) 25 (1.5 mm.) (E. A. Smith, 
1898 ) 


The radule of two alcoholic specimens were examined 
(fig. 5). The formula is C 1/1; I 37/2 + 1/1 = 38 — 1 — 38. 
The central has a single heavy cusp. The laterals have a 
broad, thin blade (mesocone), and a stout, recurved aculeate, 
smaller one (entocone). The edge of the mesoconal blade 
from the 7th out is quite, noticeably concave near the tip. The 
base of each of the teeth is narrow, elongate and poorly lim- 
ited posteriad; that of the laterals is divided longitudinally, ex- 
cept near the base, by a groove which runs up to between the 
cusps. The outermost teeth are reduced in size; the last is 
little more than a mere denticle. This radula appears closest 
to that of M. pontificus (Gould), as figured by Pilsbry (1904). 


Brachypodella raveni (“Bland” Crosse) (1872). 
Thirty-two specimens, some living ; from underside of rocks. 
CERIONID/£ 
Cerion uva (Linn.) (1758). 


Very abundant; cemented to the rocks and the lower por- 
tions of the vegetation. None of my specimens belong to the 
var. desculptum Pilsbry and Vanatta (1896). 


8 University of Michigan 
Part II. ‘TERRESTRIAI, OPERCULATES OF VENEZUELA. 


In order to escape useless repetition, the following habitat 
and station symbols (preceded by H) are used throughout this 
paper. The last number of these references is that of the 
station. Another part will describe the habitats and localities 
in greater detail. Many of these places have been describe: 
by E. B. Williamson, in number 130 of this series (1923). 
My spelling of place-names is mainly copied from the Vene- 


zuelan government maps. 


Terrestrial Habitats 


H, I. Mountain sides; heavy forest. 
H, I, a. Rock slopes and faces. 
H, 1, b. Talus and leaf mould. 
H, Lx... Rootumonuid: 
H, I, d. Leaves and stems of trees and shrubbery. 
H, Il. Lowland, and stream flats; heavy forest, 
(letters as in H, I). 
H, III. Cultivated fields. 
H, IV. Artificial savannahs. 
H, V. Second growth forests, tombas, coffee and cacao plan- 
tations (montado), 
(letters as in H, I). 


Aquatic Habitats 


H, VI. Mountain brooks (quebradas). 

H, VI, a. Waterfalls and rapids. 

Ho V1, bs . Pools. 

H, VI, c. Spring swamps, or separate pools on flats. 
H, VII. Mountain creeks (rios), 

(letters as in H, VI). 


Occasional Papers of the Museum of Zoology 9 


H, VII. Lowland or valley brooks (canos and quebradas), 
(letters as in H, V1). 

H, IX. Lowland or valley creeks (rios), 

(letters as in H, VI). 

, X. Rivers (trios). 

, XI. Large lagoons (lagunas). 

, XII. Savannah ponds (lagunas). 

, XIII. Forest pools, mainly temporary. 


XIV. Brackish water lagoon. 


, XV. Breakwater (ocean). . 


Stations 
La Guaira, Federal District. 
(Peers Oo; coast }* 
1. Valley of Rio Macuto. 


San Esteban, Estado Carabobo. 
(L 11, 67; about 100 meters altitude.) 
2. Along Rio San Esteban, near the village. 
3. Quebrada Grande; opposite and about 2 kilometers above 


the town. 


4. Ridge east of town. 

5. Ravina de las Palmas; opposite and just below the village. 
6. Sides of Cumbre Chiquito; above and opposite Las Qui- 

guas. 
Bejuma, Estado Carabobo. 
(L 10, 67; about 600 meters altitude?) 

7. Banco Largo. 

8. Rio Bejuma Valley. 


*Throughout this paper, the numbers in parentheses preceded by 
the letter (L) indicate north latitude and west longitude to the near- 
est degree. The altitudes in the list of stations are those of the 
towns. 


IO 


16. 


17: 
18. 


TQ. 


University of Michigan 


Laguna de Ramon Coronel. 

Rio Aguirre and Sabanas de Aguirre. 
Quebrada west of Banco Largo. 

Rio Bejuma at Bejuma. 

Rio La Mona (Caserio Silva). 


Rio Chirgua (near Caserio Silva). 
Cerro Chiriguara. 


Miranda, Estado Carabobo. 
(120, -68,) 
Miranda. 
Nirgua, Estado Yaracuy. (L 10, 68; 867 meters altitude). 
Wooded ridges of La Chapa (1109 meters altitude). 
Rio Pina, above intake (headwaters of Rio Nirgua). 


Rio Nirgua and savannahs near town. 


San Felipe, Estado Yaracuy. 


(1, 11, 68; 226.5 meters altitude). 


Palma Sola, Estado Yaracuy. 
(L, 11, 68; 226.5 meters altitude). 
Around town and towards Canto Minapam. 
Along Rio Aroa. 
Hills along railroad towards San Felipe. 
Aroa, Estado Yaracuy. (L 11, 68; 212 meters). 
Quebrada Carampampa (?); west of Aroa. 


Mines of Aroa and above. (Quebradas Honda and Las 
Minas). 


Boquerén, Estado Yaracuy. 
(1, 11, 68; about 125 meters altitude.) 
Rio Aroa. 


Quebrada Vaca (tributary of Quebrada Carabono). 


. Quebrada Cobre. 


Occasional Papers of the Museum of Zoology II 


27. Woods around Boqueron. 

27a. Rio Yumarito. 

28. Near Quebrada Seca, Cerritos de Yumarito. 
29. Quebrada Sucremo. 


Tucacas, Estado Falcon. 
(0.11, 68 3..coast.) 
30. Laguna near Tucacas. 
31. Around intake on Rio Tuca. 


Puerto Cabello, Estado Carabobo. 
(L 11, 67; coast.) 
32. Breakwater. 
Maracaibo, Estado Zulia. 
(L 9, 72; coast.) 
33. Brackish water lagoon in suburbs. 


Rio Catatumbo, Estado Zulia. 
(L 9, 72; coast.) 
34. Boca Norte del Rio Catatumbo, Laguna de Maracaibo. 


Estacion Tachira, Estado Tachira. 
(L 8, 72; 364 meters altitude by barometer. ) 
35. Brook, on opposite side of Rio Lobaterita, just above 
town. 
36. Quebrada Uraca. 
37. Camino Real, near mouth of Quebrada Uraca. 
38. Quebrada just west of town, and right side of cafion of 
Rio Lobaterita. 


La Fria; Estado Tachira- 
(L 8, 72; 140 meters altitude.) 
39. Cafio de las Brujas, Cerritos de las Brujas. 


40. Heavy, lowland forest, between railroad track and Rio 


ba Griz: 


12 University of Michigan 


41. Quebrada La Fria, near Camino Real. 
42. Quebrada Santa Aguita, near Camino Real. 
43. Between Quebrada Las Pipas and Rio Oropito. 


44. The “tombas” at the railroad bridge across Rio La Grita. 


El Guayabo, Estado Zulia. 
(L 8, 72; 68 meters altitude. ) 


45. Near Rio Zulia (El Cafio Fraile). 


Encontrados, Estado Zulia. 
(U0, 72; 42 meters: alitiade,) 
46. Ponds and river flats near Rio Catatumbo, northwest of 


town. 
NERITIDA 
Neritina virginea reclivata (Say). 

Seventeen specimens from slightly brackish lagoon near 
Maracaibo (F. B. Williamson). These specimens resemble 
closely those from Florida, although they somewhat approach 
N.v. microstoma from Cuba. A few show more widely-spaced 
hair-lines than is common in typical N. v. reclivata. The gen- 
eral coloration is dark olive-green to olive-brown. ‘The radula 
of specimens from this lot have been figured in another paper 
(1923, Proc. Acad. Nat. Sci., Philadelphia). Individuals 


measure :* 


altitude major diameter 
13.7 97 (13.3). 
12.2 102° (E25 
10.8 99 (10.7) 


2 Throughout this paper, the altitude of the shells is given in milli- 
meters. The other dimensions are expressed as an index, the dimen- 
sion divided by the altitude, in percentages of the latter. The dimen- 
sion index is followed by the actual measurement in millimeters (in 
parentheses ). 


Occasional Papers of the Museum of Zoology 13. 


Nerita tessellata Gmelin. 


The radula of specimens, from the break-water at Puerto 
Cabello (H, XV, 32), has been discussed in another paper 
cee.). 

HELICINIDA 


Oligyra (Alcadia Sericea) riparia tachirensis, 


new subspecies 


‘ighteen specimens; Estacion Tachira and La Fria, from 
leaves of trees and shrubs, and on ground in rich humus (aes- 
tivating), on mountain sides and in lowland jungle (H, I, bd, 
35; II, bd, 40). This species is usually found nearer the ground 
than is Helicina concentrica. : 

In the A. N. S. P. is a set (no. 14612) of three, bleached 
specimens of H. riparia Pfr. (1854), from near Calamar, Co- 
lombia (L, 10, 75, the type locality). They still show signs of 
the spiral lines and the peculiar indentation of the basal edge 
of the peristome, and appear to be quite closely related to O. 
sericea (Drouet, 1859), from Cayenne. Other forms of the 
latter species are O. sericea paraensis (Pir., 1859), from Para, 
Brazil, and O. sericea kuchm (Pfr., 1872), from Surinam. 
However, O. riparia is a larger, more depressed shell, with a 
tendency towards more definite, spiral lines, and more rapidly 
enlarging whorls. Fresh specimens, of a brown color form, 
from Aracataca, Colombia (L 11, 74), a few miles northeast 
of the type locality (A. N. S. P. no. 46579, Rehn and Hebard, 
1920), show the hairy epidermis of the group. 

O. r. tachirensis is very similar to the typical form, but is 
usually larger, higher, and with a tendency towards less marked. 
sp ral lines. Two, quite distinct, color forms occur: dull chest- 


nut brown (10 specimens) and pale lemon yellow (8 speci- 


14 University of Michigan 


mens). All of the shells are unicolor. The growth lines are 
markedly definite and regular; in some cases, spiral lines are 
also quite prominent, but this character is variable. The short 
(.5 mm.), reddish-brown hairs are often removed with the in- 
crusting dirt, but, in perfect specimens, are arranged so as to 
form spiral rows and also rows parallel to the growth-lines. 
The heavy, basal callus has a peripheral thickening, where it 
appears as a projection at the base of the columella. In young 
specimens, just outside of this projection is a definite, triangu- 
lar slit, which is represented by a well-marked indentation in 
fully mature shells with slightly reflected and thickened peri- 
stome. ‘The heavy and well-developed, calcareous plate of the 
operculum is similar to that of O. sericea (Cf. Wagner, 1907, 
X-4), but bears a more definite, longitudinal ridge on the inner 
surface, running from the nucleus to the basal angle. ‘The an- 
gular region is also especially thickened. The very thin, horny 
plate is slightly pinkish in color; during removal of the animal, 
it is usually separated from the calcareous plate, as it adheres 
closely to the foot. 


The type belongs to the brown color-form, and was collected 


on leaves of trees near La Fria (H, II, d, 4o). 


Whorls Height Maj. Diam. Min. Diam. 


O. sericea 4 5.0 120 (6.0) 100 (5.0) Wagner, (1907) 
O. riparia 5% 5-0 140 (7.0) 125 (6.2) Pfeiffer (1854) 
Sat. eeAOn 7 O15 4.8 140 (6.7) 119 (5.7) Aracataca, Col. 


O. r. tachirensis 4% 5.6 127 (7.1) 114 (6.4) Female (type). 
4% 5.0 128 (6.4) 110 (5.5) Male. 

‘The radular formula of O. r. tachirensis is given in another 
paper (1922), and characteristic teeth are figured here (fig. 
iii-14). ‘The very broad, rachidian plate distinguishes it from 
Alcadia s. s.. Other species of Oligyra from northern South 
America are: O. (Analcadia) dysoni dysoni (Pfr., 1859), 


Occasional Papers of the Museum of Zoology 15 


from Cumana, Venezuela (L 11, 64); O. dysoni barbata 
(Guppy, 1864) from Trinidad; O. (Succincta) cacaguelita 
(Pilsbry and Clapp, 1902) from Cacaguelita, Santa Marta 
Mts., Colombia (lL, 11, 74); and O. (?) pellucida (Sowerby, 
1842) from Surinam, Guiana. 

The radular formula of a dried specimen of O. dysoni bar- 
' bata (A. N.S. P. 14917, Trinidad, Swift Collection) is given 
below and samples of the teeth are figured (fig. iii-12). The 


Uncini with 


A B Cc D 3-cusps; 4-cusps Total Each row 
O. dysont barbata 3-4 5 A. 7-8 8 4 64 139 
O. rufa 3-4 5 45 7-8 6 4 75 101 


centrals are quite similar to those of Oligyra s. s., but the shell 
characters approach those of Alcadia; Analcadia is perhaps 
best retained as a subgenus between the two subgenera Oligyra 
and Alcadia. The radular formula of two dried specimens of 
O. (Analcadia) rufa (Pfr., 1857) from Yuma River, Haiti 
(A. N. $. P. 60953), is given for comparison (fig. ili-I1). 
The inner marginals have two large cusps and a vestigial, outer 
one, very much as in Idesa and Alcadia s. s. Otherwise, this 
radula is very similar to that of the South American species. 

The radular formula and probable position of O. cacaguelita 
have been discussed in a former paper (1922). I have not 
seen specimens of O. pellucida, but it also appears to belong 
in Succincta. However, other writers have noted its similarity 
to H. tamsiana Pfr. 

For comparison with the above, the radula of O. (Alcadia 
Idesa) rotunda (Orbigny, 1845) from Marianao, Cuba, is fig- 
ured (fig. 111-13). The formula is given in another paper (1922). 
It agrees with Alcadia s. s. in the shape of the centrals, but the 
pointed cusps of the D-plate resemble those of Sericea or Anal- 


cadia. ‘The first marginal is practically bicuspid. 


16 University of Michigan 


Helicina (Tristramia Oxyrhombus) concentrica Pfr. (1849), 
and approaching var. pandiensis A. J. Wagner (1905). 


Sixty-eight specimens; Estacion Tachira and La Fria, from 
leaves of trees, palms and shrubs, and from the ground (aesti- 
vating ), mainly in rich humus, but also in the root mould (H, 
Tabd 935) 42,433 LI, bd-40; 43). 

The type locality of H. concentrica concentrica is near Merida, 
Venezuela (L, 8, 71). while that of H. c. pandiensis is Pandi, 
Colombia (Ll, 4, 75). These lots, from an intermediate region, 
contain specimens that appear to be very close to both of these 
subspecies. ‘The growth lines are rather widely spaced; under 
the lens, all of my specimens also show fine, anastomosing lines, 
which tend to run spirally. In addition, a few obscure, but 
quite regular, spiral furrows are usually, but not always pres- 
ent, both on the apical and basal sides of the whorls. The 
thread carina is usually sharp. The columellar angle of the 
peristome is usually rounded, but is often quite sharply angu- 
late. The smooth apex is greenish to golden-yellow in color; 
the callus, peristome, and the edge of the carina are whitish to 
yellowish. The ground-color varies, in the series, from light, 
chestnut brown to lemon yellow ; it is often darker towards the 
carina and sometimes is broken by obscure, spiral bands or 
lines. As shown by the measurements, this species shows con- 
siderable variation in size and shape. Some of the specimens 
have a slightly scalariform tendency. ‘The specimens from 


‘stacion Tachira are small and depressed. 


Occasional Papers of the Museum of Zoology 17 


Whorls Height Maj. Diam. Min. Diam. 


H. c. concentrica 4% 6.0, . 167 (10:0) « 133. (8.0), Pfr. (1852): 

4% 6.5 138 ( 9.0) 123 (8.0) A.J. Wagner, (1905). 
6.5 139 (9.0) 115 (7.5) A.J. Wagner, (1905). 
5.9 71 (10.1) 149 (88) Estacién Tachira. 
6.4 170 (10.9) 140 (9.0) 
6.25) 250, C'O.7) Iga °(&s) La Fria: 


H. c. pandiensis 5 
5 
5 
5 
5 6.6 144 (9.5) 124 (82) 
5 
5 
5 
5 


H, I, b, 35 
El, bd, 40 


Fale y EST ((IO:7); \ c126"1(0:0) 

734259 (11-6) 238( 10-3) 

7.5 144 (10.8) 126 (9.4) 

£0), 153 (12:3) + IrCios) 
The radular formula of H. concentrica is given in another 
paper (1922), and is figured here (fig. iv-15). A surprisingly 
large number of forms, apparently belonging to this group, 


have been described from the northern Andes. 


Helicina (Tristramia Angulata) rhynchostoma ernesti von 
Martens (1873), and approaching var. infesta A. J. Wag- 
ner (1905). 

Twenty-five specimens; San Esteban, Palma Sola, and Bo- 
queron, from leaves of vegetation and in rich humus on the 
ground (aestivating), in mountain and lowland forests (H, I, 
b, 7; II, bd, 20, 27, 28). This species appears to be more 

- truly arboreal than is H. tamsiana, and is usually found farther 
from the ground. 

The shape of these specimens is quite variable. The basal 
callus is rather light, and is bounded by a distinct sulcus. The 
basal angle of the peristome is either rounded, or thickened so 
as to be quite angular. The spiral lines may be deeply im- 
pressed and numerous, or almost absent. The color is usually 
lemon yellow, with a pinkish purple band above and below the 


thread of white on the carina. 


18 University of Michigan 


Height Maj. Diam. Min. Diam. 


H. r. ernesti 10.0 180 (18.0) 140 (14.0) von Martens (1873). 
So 175 (aie) | 137 (16.0) 
H. r. infesta 11.0 164 (180) 145 (16.0) A.J. Wagner (1905). 
A.N.S.P. 14629 89 192 (17.1) 160 (14.2) Puerto Cabello. 
9.6 167 (16.0) 147 (14.1) CE. infesta. 


Eel aD 27, 7.2 200 (14.4) 165 (11.9) San Esteban. 
8:0' “19619 (i157) ” 166 (Cr3a) 

HP aE vde20 97.5 200 (15.0) 167 (12.5) 
8.5 187 (15.9) 160 (13.6) 

eit 25 7.4 185 (13.7) 160 (11.8) Boquer6n. 
8.6 107 (16.9) 166 (14.3) 


This form has often been confused with H. candeana Sow- 
erby (1842), from Honduras. After comparison of the de- 
scription and figures with the specimens before me, H. infesta 
seems little more than an extreme form-variation of H. r. 
ernestt. Other species from northern South America are: H. 
rhynchostoma rhynchostoma “Shuttleworth” Pfr. (1865) from 
Campanera, Colombia (L ???); H. ocanensis A. J. Wagner 
(1905) from Ocana, Colombia (L 8, 73); H. steindachnert 
and the cotypical form superstructa A. J. Wagner (1905) from 
Frontino, Colombia (L 7, 77); H. colombiae E. A. Smith 
(1878) from San Sebastian, New Granada (L 11, 747); and 
H. gonochila Pfr. (1849) (+-gonocheila Sowerby, 1874) from 
Venezuela (Caracas?). I have seen no authentic specimens of 
any of them, but, from descriptions and figures, the second ap- 
pears to be quite distinct, the third rather close to H. rhyncho- 
stoma; the fourth is unfigured, but seems to differ from the 
first by its smaller size and higher spire. H. gonochila closely 
resembles H. braziliensis Gray (1824), and its Venezuelan hab- 
itat appears open to doubt. 

The radular formulae of H. r. ernesti and H. caracolla are 
given in another paper (1922); examples of the teeth are fig- 
ured here (figs. iv-16, 18). 


Occasional Papers of the Museum of Zoology 19 


Helicina (Tristramia Tamsiana) tamsiana Pir. (1850), and 


approaching var. appunt von Martens (1873). 


Eighty-two specimens; San Esteban, Palma Sola, Aroa, and 
Boqueron, from leaves of vegetation and from ground (aesti- 
vating), usually in rich leaf mould, in the mountain and low- 
iid torestsaCm, J.-d0, 2, 5.23 lL bd, 20, 22, 28). 

The sculpture of this species consists of fine, regular, close- 
ly-spaced growth-lines, crossed by numerous, fine, spiral lines, 
which are usually very regular in arrangement. ‘The consid- 


erable color variation may be analyzed as follows: 
a. Apex: golden yellow to creamy white. 
b. Peristome: white. 
c. Basal callus: whitish. 


d. Ground color: (1) fulvous, varying to reddish orange 
(typical form); (2) greenish yellow; (3) creamy white. 
Stripes are absent with the first class of ground color, present 


or absent with the second, and present with the third. 


e. Pattern: (1) Darker shading on apical side of whorls; 
when brown, this may form a distinct, dark band or be light 


and diffuse; when greenish yellow, it is always the latter. 


(2) The band of (1) may be narrowed by the presence of 
a creamy white band, just below the suture (very common 


form). 


(3) A slight infuscation may be present on the base, so as 
to mark off a narrow, light band around the greatest convexity 
of the last whorl. 


The relative numbers of the different color forms vary with 
the locality. In one or two places, all of the shells collected 
were of a single form, but, in other localities, all of the kinds 
occurred together. 


20 University of Michigan 


Height Maj. Diam. Min. Diam. 


H. t. tamsiana 6.0 150 (90) 129 (7.7) Pfeiffer (1852). 
7-5 120 ( 9.0) 103 (7.7) A.J. Wagner (1907). 


H. t. appuni 11.0 127 (14.0) von Martens (1873). 
Eek “p43 68 122 ( 83) 102 (6.9) Aroa. 
Hy ird: 20 Vfl iz Guenet), 96 (6.8) Palma Sola. 


82 213 0:95) “F96.¢7.9) 
9.4 I13 (106) 99 (9.3) 


BE IE gist, i 79 120.92) #103) (7a) + San Esteban: 
8.6 120 (10.3) 103 (8.9) 
elapse 9.4 105 ( 90) 94 (88) San Esteban. 


While these specimens vary considerably in size, none are 
as large as typical H. t. appuni (type locality given as Puerto 
Cabello). The undulation of the basal wall of the peristome, 


Figure 20. Helicina (Tamsiana) tamsiana. Centrals, laterals and 
tip of 3rd uncinus. The scale indicates a length of 50 microns. 


which caused Wagner (1907) to include this group in Alcadia, 
is seldom distinct, and usually very obscure. I have not seen 
specimens of H. columbiana Philippi (1847) from Colombia, 
but suspect that the larger H. sanctaemarthae Pilsbry and 
Clapp (1902) from Libano, Santa Marta Mts., Colombia (1. 


11, 74), is closely related. The last is more nearly conical than 


Occasional Papers of the Museum of Zoology 21 


tamsiana, has spiral rows of punctations instead of solid lines, 
and the calcareous plate of its operculum is also much heavier 
and more opaque. H. nemoralis Guppy (1866) (+ H. sonata 
Guppy, 1868, not Lesson) from Trinidad, also belongs in this 
group. The radular formulae of H. tamsiana and H. nemoralis 
are given in another paper (1922); teeth of the former are 
shown in the text figure (fig. 20). 


H. kieneri Pfr. (1849) was originally described without local- 
ity; in the Conch. Cab., it was ascribed to Guadeloupe; and, in 
1852, to Caracas. From the descriptions and figures, it seems 
to be a form of the protean H. rhodostoma Gray (1825) from 
Guadeloupe. In this connection, it may be remarked that there 
is a group of Caracas Islands off the coast of Cumana (L Io, 
65). H. crassilabris Philippi (1847) is undoubtedly a Pacific 
species, probably from Sandwich Island, New Hebrides. 


Lucidella (Lindsleya?) venezuelensis, new species 


Twelve specimens from damp leaf-mould in a hollow left by 
a temporary forest pool, near Cafio Minapam, Palma Sola (H, 
Xill, 20). 

Shell: minute, conic-globose, perforate. Color: whitish horn, 
tinged with orange towards the apex. Whorls: 5%, convex; 
apical 34 whorl, minutely raised-punctuate; the remainder with 
fine, closely-spaced, spiral riblets, which extend on to the base 
of the shell; last whorl tapers toward aperture and is free 
(solute) for a short distance. Aperture: kidney-shaped, with 
oblique, almost straight, parietal wall; upper angle rounded. 
Peristome; free, simple, and very slightly expanded. Oper- 
culum: concentric, with calcareous plate thicker than the horny 
one, and markedly concave externally; spiral nucleus almost 
central; horny plate almost flat, so that its edge is markedly 
separate from the calcareous one. 


22 University of Michigan 


The type (fig. ii-7) measures: 
_ Altitude Diam. Length Width 

Height Maj.Diam. Aperture Aperture Operculum Operculum?® 

3.05 93 (2.85) 38 (1.15) 104 (1.20) 46 (1.40) 72 (1.00) 

The systematic position of this species has been discussed in 
an earlier paper (1922), where a preliminary description of 
the radula i8 given. The figure of the teeth, included here 
(fig. iv-17), shows its close relationship to Lucidella. The 
shell is certainly closest to that of Stoastoma domingensis Van- 
atta (1920), but it is slightly larger, more depressed, and has 
a larger number of finer, spiral riblets than the species from 
Santo Domingo. 


Lucidella (Poema) lirata (Pfr., 1847). 


Widely distributed, but especially abundant in damp and in 
disturbed places: San Esteban, Palma Sola, Aroa, Estacion 
Tachira, and La Fria; from damp, rich humus on mountain 
slopes, in lowland forests and cacao plantations, in a spring 
swamp, and in the damp depressions occupied by forest poois 
during the wet seasons (H, Ib, 2, 5, 23, 35; U1, b, 405 Vag 
2, 41; VI, c, 38; XIII, 20). These specimens are mainly quite 
characteristic of L. lirata lirata, but a few show some approach 
to L. lirata lamellosa (Guppy, 1867) from Trinidad. 

For comparison, the radular formula (fig. iv-19) of two 
dried specimens (A. N. S$. P. 62062, St. Kitts, W. H. Rush, 
1891) of L. plicatula christophori Pilsbry (1897) is given: 


R; At/1; B1/6-7; C1/§; D1/o; B; M11/3+ 9 (42/44+-)=53; (117). 


3 The altitude and diameter of the aperture are measured parallel 
to the long and transverse axes of the shell. The length and width 
of the operculum are measured along its long axis and at right an- 
gles to it. The index of the diameter of the aperture is that dimen- 
sion divided by the altitude of the aperture. The index of the width 
of the operculum is the width divided by the length. 


Occasional Papers of the Museum of Zoology a 


The radula of this species is quite close to that of Poenia s.s., 
but the shell sculpture consists of axial, instead of spirai rib- 
lets. This seems a valid hasis for the separation of a new 
section: Poeniella, type H. (plicatula) christophori Pils- 
bry (1897) from St. Kitts. L. ignicoma (Guppy, 1868) from 
Trinidad, appears to be a member of this group. 

In addition to the above Helicininae, Eutrochatella (sub- 
genus s.s.) semilirata (Pfr., 1849) has been described from 
Venezuela. As it has not been reported since, the presence of 
this subgenus in South America still remains very doubtful, 
although a member of the subgenus Pyrgodomus might be 


expected. 


PROSERPINID-= 
Proserpina (Linidiella) swifti Bland (1863). 


One specimen, without animal, from leaf-mould in corner 
of buttressed tree, along Rio San Esteban (H, I, b, 2). It 


measures: height 4.6 mm.; maj. diam. 187 (8.6 mm.). 


POMATIASID 
Tudora plicatula (Pfeiffer). 


Cyclostoma plicatulum Pfeiffer (1846). Puerto Cabello, Venezuela. 
?’Cyclostoma tamsiana Pfeiffer (1850). Puerto Cabello. 

?Cyclostoma cumanense Pfeiffer (1851). Cumana, Venezuela (LL 11, 64). 
?Cyclostoma venesuclense Pfeiffer (1853). Venezuela. 


Ninety-six specimens; mainly from rather well-drained for- 
est (H, II); at Palma Sola (stations 20, 22), and Boqueron 
(stations 27, 28); the most abundant large snail, wherever 
found. This species aestivates under leaves in rich leaf- 
humus (H, II, b), but, during rains, soon travels up into the 
vegetation (H, II, d), as high as 3 meters above the ground. 


Its jerky movements are more rapid than those of most snails, 


24 University of Michigan 


and it snaps quickly into its shell when disturbed. It occasion- 
ally suspends itself from leaves, by a mucous thread 2 or 3 cen- 


timeters in length. 


The operculum of T. plicatula has been described as like 
that of Chondropoma (Cf. Henderson and Bartsch, 1920), 
but fresh specimens have a well-developed, calcareous portion, 
parallel to the “‘chondroid” plate, and the outer, calcareous 
plate is supported by fine, vertical lamellae parallel to the 
growth lines. Each whorl of the calcareous portion is not 
quite so wide as the underlying whorl of the horny, basal plate. 
On account of its thin, fragile nature, the calcareous plate ts 
quite easily crushed and brushed away, so that only the chon- 
droid portion is left. Its fragility may be correlated, in part, 
with the acidity of the leaf-mould, and the absence of lime- 
stone in the region studied. Structurally, this operculum does 
not differ greatly from that of T. williamsoni (see below), but 
its whorls increase more rapidly in size, so that the nucleus ~ 


is more eccentric. (See fig. 11-9.) 


The shells are very variable in color and texture. Usually, 
spiral bands (as many as 10), of chestnut brown to purplish, 
are present, but these are almost always broken into dots. The 
dots tend to be arranged in vertical rows and, in a few speci- 
mens, these are joined together by dark varices, so that the 
bands become vertical instead of spiral. The ground-color 
varies from light cream to dark fulvous. Most of the speci- 
mens from the edge of the hills (stations 22, 28) are dark 
colored, and are very commonly quite unicolor. 

The sculpture varies from rounded ridges parallel to the 
growth lines, to marked, thread-like riblets. Usually some 
buttress-like denticulations are present in the sutures, but their 


number and size is very variable. The strength of the umbil- 


Occasional Papers of the Museum of Zoology 25 


ical spiral is also variable, but, in all of my specimens, at least 
two can be detected. In addition, traces of spirals are present 
on the sides of the earlier whorls. The aperture of adult spec- 
imens is always solute, angulate above, and with the peristome 
free and slightly expanded. This expansion is most noticeable 
on the lower portion of the palatal wall, but is least extensive 
along the base. 

The females are considerably larger than the males: repre- 
sentative examples of each measure: 


Altitude Maj. Diam. Altitude Aperture Diam. Aperture 


Female 17.3 53 (9.2) 38 (6.5) 85 (5.5) 
Males 14.5 54 (78) 36 (5.2) 87 (4.5) 
12.4 55 (6.8) 34 (4.2) gr (3.8) 


Pfeiffer also described Cyclostoma tamsiana from Puerto 
Cabello. According to his description, the calcareous plate of 
the operculum is the only important character that separates it 
from C. plicatulum. As detailed above, this character is a 
doubtful one. However, he emphasizes the central position 
of the nucleus of the operculum of C. tamsiana, and it is just 
possible that he had immature specimens (peristome simplex) 
of T. williamsont secana (see below) before him. Pfeiffer’s 
own figure (Conch. Cab., xxxvii-19, 20) might represent the 
latter, but Reeve’s (1863, Conch. Icon.) is certainly T. plica- 
tula, and Pfeiffer (1865) recognized it as authentic. All of 
the Puerto Cabello specimens in the A. N. S. P. are T. plica- 
tula, although some of the lots, with better preserved opercula, 
have been labeled tamsiana. In this connection, attention 
should be called to the habit of earlier collectors, who very 
commonly named, as the locality of their specimens, some 
large, well-known town in the general region. Labels such as 
Puerto Cabello and Caracas may mean but little more than 


labels like Venezuela or New Granada. 


26 University of Michigan 


I*rom the variation in color and sculpture of my specimens, 
I very much question the specific rank of. C. cumanense and 
C. venezuelense, but have seen no authentic examples. Ch? 
subauriculatum Pfr. (1862) was also described from Cumana 
(Bland). Specimens from Bland (original lot?) in the A. N. 
S. P., show this to be a heavier form near C. biforme Pfr. 


(1858), from Turks Island, Bahamas. Bland’s locality is 
probably incorrect. 


Tudora williamsoni, new species 


Eighty-five specimens from damp, bare, rock faces along 
quebradas in Aroa Mountains (H, I, a, 23 (type), 24). The 
most abundant shell present in these places, although Brachy- 


podella hanleyana is almost as common. 


The shape of these shells is very similar to that of T. plica- 
tula, but the 614 to 634 whorls are more convex, and the su- 
tures more deeply impressed. The shells are heavier; the 
growth riblets are high, but compressed, and tend to occur in 
pairs, or more rarely in groups of 3 or 4. The buttress-like, 
sutural prolongations of the riblets are fine, but prominent. 
The spiral, umbilical sculpture is present, but is obscured by 
the crowded riblets (fig. 11-8). The last whorl is free (so- 
lute) for a very short distance. The aperture is almost circu- 
lar. The peristome is quite broadly (.8 mm.) reflected on all 
sides, but most prominently on the columellar lip, which is 
often undulate. The parietal portion is produced into an an- 
gular auriculation, which is adnate to the preceding whorl. In 
adult shells, the reflection consists of several layers of varying 
extent, so that a polyplex condition is produced. The color- 
markings are similar to those of T. plicatula, but the higher 


riblets give the shell a duller appearance. 


Occasional Papers of the Museum of Zoology 27 


The structure of the operculum (fig. ii-8) is quite similar 
to that of T. plicatula (fig. ii-g), but the calcareous plate is 
heavier and more extensive, and is less apt to disintegrate in 
the dried specimens. ‘The whorls of the operculum increase in 
size very gradually, so that the outline is almost circular, and 
the nucleus practically central. Its appearance is almost as 
close to that of Annularia as to that of Tudora megacheilos 
(fig. 11-10). 

The general appearance of this species is closest to that of 
T. aripensis (Guppy, 1864) from Trinidad. The latter has 
more regularly spaced riblets and the shape of its operculum 
is more or less intermediate between that of T. plicatula and 
that of T. williamsoni. 

Specimens measure: 


Height inside Diam. inside 
Altitude Maj.Diam. of Aperture of Aperture 


Largest female 12.1 55 (6.6) 261 (3) 100 (3.2) 
Another. female 10.6 57 (6.0) i (PATH) 96 (2.6) 
Male; type TRE: 67 (4.8) 33)4(2:4) 100 (2.4) 


Tudora williamsoni secana, new subspecies 


Eleven specimens; on rocks and bare clay banks of an arroya 
in the Cerritos de Yumarito, near Quebrada Seca (H, II, a, 
28). With it occurred Brachypodella hanleyana, and, in the 
leaf humus nearby, Tudora plicatula. 

This subspecies differs, from the typical form, mainly in its 
larger size. In addition, it is usually heavier, with darker 
ground color and more obscure color pattern. The males of 
this lowland form are of about the same size as the females 


of the mountain one. Representative specimens measure: 


Height inside Diam. inside 

Altitude Maj.Diam. of Aperture of Aperture 
Female (type) 14.1 54 (7.6) 201 (3:7) 100 (3.7) 
Male 10.7 62 (6.6) 25 (2.7) 100 (2.7) 


28 University of Michigan 
CYCLOPHORIDAE 
APEROSTOMATINAE 


The synonymy of the major groups of this subfamily has 
been reviewed in an earlier paper.* Of the South American 
genera, Buckleyia appears to extend only as far north as south- 
ern Colombia. Besides the numerous species of Amphicyclotus 
from Central America and northwestern South America, A. 
cayennensis (Shuttleworth, 1852), from Cayenne, Guiana, ap- 
pears to belong to the same group as the species from the Les- 
ser Antilles. With the exception of the doubtful Cyclostoma 
psilomitum (see below), the subgenus Neocyclotus, of the gen- 
us Poteria, is the only group that has been reported from Ven- 


ezuela. 


A brief synopsis of the mainland species of Neocyclotus will 
be presented. All of the species included here, with the ex- 
ception of P. corpulenta (Smith), have been figured. The 
most recent bibliographies are found in Kobelt (1902, Thier- 
reich) and Kobelt-Schwanheim (1912, Conch. Cab.). In the 
following synopsis, those species preceded by an exclamation 
point (!) are not included in either of these monographs, but 
are figured by their authors. Those preceded by an asterisk 
(*) are not in the Conch. Cab. 


Although Pfeiffer (Conch. Cab.) describes the operculum 
as calcareous, I very much doubt the position of Cyclostoma 
prominula “Ferussac” Orbigny (1835) (+ Cyclostoma bra- 
siliense Sowerby, 184—, + C. planorbulum Sowerby, 1842), 
as a member of the subgenus Neocyclotus. The apical whorls 
are those of the Lesser Antillean group of Amphicyclotus, 
while the solute last whorl is quite distinctive, although it oc- 


* “Aperostomatinae,” The Nautilus, xxxv, July, 1922. 


Occasional Papers of the Museum of Zoology 29 


curs as an abnormality in other species. ‘The type locality is 
Minas Geraes, Brazil. It is just possible that this species is. 
related to Cyclostoma disjunctum Moricand (1846), also from 
Brazil.* 

The opercula of (*) Cyclostoma psilomitum Pfr. (1851) 
from Venezuela, Aperostoma connivens H. Adams (1866) 
from Eastern Peru, and Cyclostoma distinctum Sowerby 
(1843) from the Bay of Montija, Panama, are unknown, but 
these Crocidopoma-like species certainly do not belong in Neo- 
cyclotus. (*) Cyclotus coopert Tryon (1863) from Mazatlan, 
Mexico, is clearly an Amphicyclotus, close to A. lutescens 
(Pfr., 1851); while (*) Cyclotus boucardi Angas (1878) 
from San Carlos, Costa Rica, not A. boucardi (Pfr., 1856) is 
evidently close to A. ponderosus (Pfr., 1851). (!) N. inflatus, 
from Santa Marta, and (!) N. solutus, from New Granada, 
both “Mousson” Kobelt and Moellendorff (1897, Nach. D. 
Malac. Ges.) appear to be nude names, as is also (!) N. 
chrysacine “Bartsch” Fluck (1906, Naut.), from Wani, Nic- 
aragua. Cyclostoma suturale and C. discoideum, both Sower- 
by (1843), are extralimital species, incorrectly cited from 
Demerara, Guiana. 

The mainland species of the subgenus Neocyclotus may be 


divided into four, quite distinct groups, as follows: 


1. Group of Cyclostoma translucidum Sowerby (1843). 
Shell turbinate, usually unicolor; growth lines very prominent 
and often crossed by spiral striations; basal portion of peri- 
stome usually slightly emarginate. Mexico to Trinidad and 


Peru. 

5 C, disjunctum is the type of Cyclopoma Troschel (1847), not Agas- 
siz (1833); this preoccupied name was inadvertently omitted from my- 
1922 paper. It is a synonym of Amphicyclotus. 


30 University of Michigan 


2. Group of Cyclostoma stramineum Reeve (1842). Shell 
turbinate or depressed, usually unicolor; growth lines crossed 
by oblique grooves and plicae, which break the sculpture into 
rugae or granulations. Ecuador (?); Panama to Lesser An- 
tilles. 


3. Group of Cyclostoma inca Orbigny (1835). Shell de- 
pressed conoid, or with last whorl descending markedly (Cf. 
Poteria confusa); usually with darker, spiral bands; growth- 
lines comparatively delicate and regular. Central America to 


Brazil. 


4. Group of Cyclostoma incomptum Sowerby (1849) from 
Brazil. Shell similar to that of group 3, but with siphon-like 
development of the upper edge of the peristome. Colombia to 
Brazil. N. pergrandis Kobelt-Schwanheim (1912), from 
“New Granada,” is a larger shell than the typical species. 


In the localities studied, the forms of Poteria were found 
to be remarkably limited in their distribution. In only one 
place did two species occur together, and there the mixture 
occurred only at the junction of two distinct habitats (H, I, 
II, 42). In other localities, they showed a distinct tendency 
to form small colonies. For instance, in all of the collections 
made around Palma Sola, specimens of this genus were found 
only in a single locality, and there appeared to be restricted to 
an area about 20 meters across. The general appearance of 
this area was the same as in many other portions of the sur- 
rounding forest, which makes it especially difficult to account 
for this peculiar localization. In all places, they were the 
only snails that were common in the root humus (H, I, II, c); 
- they seemed to favor especially the corners between the radiat- 


ing roots and buttresses of the larger trees. 


Occasional Papers cf the Museum of Zoology at 


Potcria translucida translucida (Sowerby, 1843). 


One almost adult shell and 5 young specimens from near 
the Rio Oropito, at the edge of the Cerritos de las Brujas 
(H, I, be, 39), appear to be quite close to this typical form. 
They are light-colored, comparatively fine-textured shells. 
Measurements of similar shells from Cucuta, Colombia (I, 8, 
73; A. N. S. P. 12963), from Puerto Cabello, Venezuela 
Nese n2eor: > Starke) hand ‘from La! Guaira, 
Venezuela (A. N. S: P. 12986) are also given (see fig. v-A). 
This subspecies appears to be characteristic of the lowlands; 
in-the mountains, the species is represented by various, close- 
ly related forms. P. translucida trinitensis (Guppy) has sim- 
ilar dimensions, but most of the specimens can be separated 
by the key (see below). The measurements of a characteristic 
example of each sex (see fig. v-B, C) from Cariaquito, Vene- 
zuela (L, 10, 63; A. N. S. P. 104631; Bond Expedition, rg1r), 
are given. In all of the subspecies, the adults examined have 


close to 41% whorls. 


Dimensions of Poteria translucida 


Height Maj Diam. 


P. t. translucida 16.0 127 (20.4) Figure, Thesaurus. 
Young shell 12.2 121 (14.8) H,I,c, 39; La Fria, Ven. 
Pees. P. 12063 12.5 131 (16.5) Cucuta, Colombia. 

12G61 12.8 127 (16.2) Puerto Cabello, Ven. 


14.5 130 (18.8) 
12986 (Fig. v-A) 16.0 133 (21.2) La Guaira, Ven: 
P. t. trinitensis (Figs. v-B,C) 13.8 130 (17.9) Male; Cariaquito, ‘Ven. 
Wee: o. PP: 104631 16.9 122 (20.5) Female; ditto. 
P. t. bejumensis (Fig. v-D) 17.2 105 (18.1) H,I,b,7; type; Bejuma, Ven. 
14.8 nL7 (zee) 
(Fig. v-E) 14.3 128 (18.3) 
(Fig. v-F) 12.0 118 (14.2) 
(Fig.v-G) 17.3 118 (20.4) H,I,c, 15; Cerro Chiriguara. 
Means: 34 specimens 14.9 rr7 (ies 
Extremes: ditto 12.0-17.3 105-128 (14.2-20.4) 


32 Umicrsity of Michigan 


Height Maj Diam. 
P.t.major; AN.S.P.12990 = 21.3 116 (26.3) Type; Caracas, Ven. (Fig. v-H), 


P.t. santaguitensis (Fig. v-J) 12.3 136 (16.7) H,1,b, 42; smallest male. 
(Fig. v-I) 178 128 (22.7) Type; largest female. 
A.N. S. P. 12993 19.2 126 (24.2) Cucuta, Colombia (Bouquet). 


The following artificial key to the mainland forms of this 
typical group may assist in their separation, although a com- 
parison of authentic specimens, or at least a study of the orig- 
inal figures and descriptions, is the only practicable means for 
their identification. The localities given are those of the orig- 
inal descriptions, except in a few cases where the original 
locality was omitted or was later shown to be absolutely incor- 
rect. The figures give the altitude in millimeters, the major 
diameter index in percentages (followed by the actual dimen- 
sion in millimeters), and the number of whorls. As the earlier 
writer determined the altitude in a different manner from that 
at present customary, many of these measurements are not 
those of the author, but are those of his figure. In a very 
few cases, the measurements are necessarily taken from later 
authors, but such cases were checked up with original data. 
The names given are those of the original authorities, and I 
do not vouch for the specific separation of many of them. As 
will be detailed later, the individual variation is often very 
great, even in shells from a single locality. 


A. Costa around umbilicus; operculum unknown. Eastern Peru. 
TAS MGO(ZE) iS Orc. cis «ee Aperostoma bartletti H. Adams (1860): 


AA. Sides of umbilicus evenly rounded. 


B. Peristome emarginate above and below; shell large. Costa 
Rica. 31; 124(38.5); 5...Cyclotus bisinuatus Martens (1864). 


BB. Peristome not emarginate above; shell smaller. 


C. Shell with comparatively weak, quite regular, growth lines; 
color light greenish to yellowish, translucid. 


Occasional Papers of the Museum of Zoology 33 


D. Shell more depressed; sutures distinctly impressed; 
whorls convex. Coloras Island, Gulf of Paria, Trin- 
adess PIGG AA RIOR OP Oe. eee ao Soe eens Se dene mes 
Soph Ee PPR 2 ee Cyclostoma trinitensis Guppy (1864). 


DD. Shell medium; sutures less distinct; whorls less round- 
eds GC oloni pina TOs T2720 Kt or 3. cn ca aeons 
ick Viale Ae Cyclostoma translucidum Sowerby (1843). 
DDD. Shell more elevated; heavier. 


E. Smaller. Bejuma, Venezuela. 14.9; 117(17.5) 4%. 

....(!) P..translucida bejumensis, new subspecies. 

EE. Larger. Caracas, Venezuela. 21.3; 116(26.3); 4%. 

Cpt she (!) P. translucida, var. major, Bland MSS. 

CC. Growth lines prominent, usually undulate; shell heavier; 

adults darker in color. 

F. Growth lines with epidermal ridgelets, usually simply 
undulate; color of adults pronounced. 

G. Color of adults dark olive green; lower lip with 

slight emargination. La Fria, Venezuela. 17.8; 128 

a) POR. ER AMES - ed oe Re oe See ee 

..(!) P. translucida santaguitensis, new subspecies. 

GG. Color of adults dark reddish; lower lip quite entire. 

‘El Libano, 6500 ft., Santa Marta Mts., Col. (L 11, 

PAN EARS S TSS CIO: ) Seater eres s,s (!) Aper- 

ostoma smithit Pils. and Clapp (1902) Nautilus. 


FF. Growth lines usually anastamosing, so as to approach 


the next group (P. straminea) in sculpture.......... 
CS ees ae bas 2 3 Poteria dysoni ( Pfr.) 
H. Peristome double, solute. Mirador, Mexico. 16; 

THOUS Go. ks. Cyclotus berendti Pfr. (1861). 


HH. Peristome simple, but often slightly solute. 

Y. Later whorls with spiral stripes of lighter color. 
Cave at Tabi, Yucatan. ..(!) C. dysoni (form) 
multiiineatus Pils. (1891). Proc. Acad. Nat. 
Sci., Philadelphia. 

II. Unicolor. 

J. Smaller. Tabi, Yucatan. Maj. diam. 16 mm. 
(!) Cyclotus dysoni, var. minor Mart. (1890).° 


® Biol. C, A.; plate 1. This is apparently a description and not a 


nhame, 


34 University of Michigan 


Jj; Larser 
K. More depressed; color castaneous. Bugaba, South Panama. 
rIsyeryan e323 (7703) IA ae Cyclotus dysoni, var. affinis Mart. (1890). 


KK. More depressed; growth lines more regular. Soledad, Mexico. 
16; 144(23).....Cyclotus dysoni, var. ambiguus Mart. (1890). 


KKK: Everything else. Honduras. 21-3); 125(26:5))s) diZ:6. 45.2522 ee 
SOR GER oo ans i. SEs Cyclostoma dysoni Pir. (1851). 


Poteria translucida bejumensis, new subspecies 
Figs. v-D, E, F, G. 

Thirty-three adults and 50 immature specimens from rock- 
talus on ledge near top of small, rock wall of cafon-like valley 
of Banco Largo, near Bejuma, Venezuela (H, I, b, 7); and 1 
adult (the largest) from leached-out, root humus near top of 
Cerro” Chirtouata (Hi, 1;c,- 45). 

Shell: heavier and more turbinate than P. ¢. translucida; 
growth lines prominent, but fine, regular and crowded. Color 
of epidermis: unicolor, yellowish horn, apex lighter. Shell 
substance: chalky, except near apex (usually eroded), where 
it is closer in texture and often tinged with rosaceous. Whorls: 
a little less than 41% ; convex, quite rapidly increasing in diam- 
eter. Sutures: prominent, but not impressed. Aperture: al- 
most vertical; circular except for upper angle; sometimes 
briefly solute. Basal lip of peristome: with one or two emar- 
ginations. Umbilicus: 1/7 of major diameter. 

The young shells, as is quite usual in the genus, are more 
depressed than the adults. The females are slightly larger 
than the males, and their average major diameter index is 
slightly greater (more depressed shell), but the individual vari- 
ation is so great that the sexes cannot be separated by their 
shape and size. 

Pfeiffer’s figure in the Conch. Cab. resembles this subspecies 


more than it does the original figure of P. t. translucida, in the 


Occasional Papers of the Museum of Zoology 35 


Thesaurus. A set of specimens (A. N. S. P. 12990), labeled 
Caracas, Venezuela (L, 11, 67), are very similar in shape to 
P. t. bejumensis, but their considerably larger size seems to 
indicate at least racial significance. They are labeled var. 
Major in Thomas Bland’s handwriting, so, with the kind 
permission of Dr. Pilsbry, that name is published here (see 
key, and fig. v-H). 


Poteria translucida santaguitensis, new subspecies 
Figs. v-I, J. 

Four adults and Io younger specimens, from root-humus 
around bases of trees on the steep valley sides of Quebrada 
Santa Aguita (H, I, c, 42), near La Fria, Venezuela. 

Shell: heavier than P. t. translucida, and growth lines ren- 
dered more prominent by fine, undulating, epidermal riblets. 
Color of adults: olive-green, translucid. Apex: tinged with 
rose. Last whorl: impressed just below suture, and often with 
a few, weak, spiral striae. Basal lip of peristome: with a def- 
inite emargination, as seems characteristic.of P. translucida. 
Umbilicus: 1/7 of the major diameter. 

The immature specimens of this subspecies are often yel- 
lowish in color. In many respects, the adults resemble P. 
smithi (Pilsbry and Clapp, Nautilus, 1902), as indicated in 
the key. A bleached specimen in the A. N. S. P. (no. 12993), 
from Cucuta, Colombia (L. 8, 73), probably also belongs to 


P. t. santaguitensts. 


Poteria straminea (Reeve, 1843). Figs. v-N, O. 


Thirty adults and 48 immature specimens, from root-humus 
on the hill-sides and around the bases of trees, in the San 


Esteban valley (H, I, c, 2, 3, 4). 


36 University of Michigan 


The inside of the aperture of these greenish-yellow shells 
is pearly, but without dark iridescence. The peristome is often 
noticeably thickened, and even flares outward, with several, 
heavy varices, so as to be almost campanulate. Its basal lip is 
quite even, and the aperture is almost circular. The exterior 
surface of the operculum is quite markedly concave, and the 
internal boss is prominent. The specimens have between 4% 
and 5 whorls, and the umbilicus is a little over 14 the major 
diameter. 

A set of specimens of Poteria granadensis rugata (Guppy), 
from Cariaquito, Venezuela (A. N. S. P. no. 104625; S. 
Brown, 1911) show that this Trinidad shell also reaches the 
mainland. Measurements of seven males and seven females 


are summarized in the accompanying table (cf. figs. v-P, Q). 


Dimensions in Group of P. straminea 
Altitude Maj. Diam. 


P. straminea 057, 142 (22.3) Figure, Thesaurus. 
Means; 30 specimens 125 147 (18.3) H, I, c, 2, 3,4; San Esteban. 
Extremes; ditto 9.7-15.9 131-158 (14.7-22.6) Figs. v-N, O. 
P. granadensis rugata 10.2(sic) (17.8) Guppy, (1864). 
A.N.S. P. 104625. 
Means: 7 females 8.6 146 (12.5) Cariaquito, ‘Venezuela. 
Extremes: ditto 8.3-9.2 138-156 (11.9-13.4) Fig. v-P. 
Means: 7 males 7.8 TA24( 17-1) 
Extremes: ditto 7.3-8.2 132-148 (10.2-11.4) Fig. v-Q. 
P. glaucostema 12.8 142 (11.4) Figure, Reeve, Conch. Icon. 
P. g. aulari 
Means: 4 specimens 15.5 146 (22.7) H, II, be, 20; Palma Sola. 
Extremes: ditto 14.1-16.8 142-154 (20.8-24.6) Fig. v-M. 
Means: 6 specimens 14.6 154 (22.6) H, II, be, 28; Quebrada Seca. 


Extremes: ditto 13.3-16.2 149-161 (21.3-24.8) Figs. v-K, L. 


The following, artificial key, to the members of the group 
of P. straminea, is presented. ‘The remarks, which precede the 
key to the first group, also apply here. The first species is in- 


cluded with considerable doubt. 


Occasional Papers of the Museum of Zoology a7 


A. Shell with irregular granulations; last whorl whitish with dark, 
chestnut-brown, spiral band. Ecuador. 18; 139(25); 5™%....... 
Cyclotus granulatus Pfr. (1862). 


Cle 6 ew 68 ©, 8 si 6's 6.6 0] oS) 6ie) 6 61 6 pj «) «6 © 


AA. Shell with comparatively regular, impressed lines, which cross the 
growth lines obliquely in two series, so as to mark off rugae or 
diamond-shaped corrugations; without definite color bands. 


B. Large species: granulations arranged in quincunx; apex liglit 
brown; lower whorls darker; unevenly black-striped. Ale- 


inno Colmmeia (1,6 76). ~ 30% 132(46.5) 3 5.035.652 0-2-3 
Neocyclotus peilei Gude (1912). 


BB. Smaller species; more depressed. 


C. Shells medium in size; plicae and striae comparatively reg- 
ular and covering practically all of last whorl. 


D. Very prominent and quite regular striae meet the suture 
at an acute angle, while the short, curved series, that 
cross the first at approximately right angles, are pres- 
ent only near the suture; color yellowish; interior of 
aperture only slightly iridescent. Merida, Venezuela 


(Tee Bieri hege Dao Gl EAS (22.9) tale iac isl. x ohhtciat «bth bi ool ale 
oor aaiocraceer teres Cyclostoma stramineum Reeve (1843). 


DD. Both series of striae comparatively weak, but almost 
equally prominent; color olive-brown to castaneous, 
lighter below; interior of aperture with bluish to pur- 
plish iridescence. 


E. Smaller; color castaneous. Venezuela. 12.8; 142 
(18.2) ; 4.(*) Cyclostoma glaucostomum Pfr.(1855). 


EE. Larger; color olive-brown. Quebrada Seca, Vene- 
REI. FAO 554 (2a ie ee ote «als wish arnaie mvs 
(!) P. glaucostoma aulari, new subspecies. 


CC. Shells smallest; both sets of striae irregular, and obscure 
or absent on lower side of last whorl. 


F. “Obscure retuse corrugati’; golden unicolor. Chiriqui, 
Pagani. A072 TSOCIS) } AVGer amen ow os) vase sane avade 
sgh hits At BS ed Neocyclotus panamensis DaCosta (1904). 


FF. Sutural side of last whorl with irregular, but promi- 
nent, striations and rugae; umbilical side obscurely 
granulate or without oblique striations; color yellow- 
ish. Northern Hills of Trinidad. (Other similar 
forms in Lesser Antilles; compare Cyclostoma grana- 
dense Shuttleworth, 1857). Maj. diam. 17.8.......... 
PPPS CUR vad ea gaces Cyclotus rugatus Guppy (1864). 


38 University of Michigan 


Poteria glaucostoma aulari, new subspecies 
Figs. v-K, L, M. 


Six adults and 12 immature specimens from the edge of 
the Cerritos de Yumarito, near Quebrada Seca, Estado Yara- 
cuy, Venezuela (H, II, be, 28; type locality), and 4 adults 
and 3 young from small colony in heavy forest, 2 kilometers 
south of Palma Sola (H, II, be, 20). 


Shell: large, but thin. Color: above, dark olive-brown with 
chestnut tinge; below, olive-green, the line of demarcation 
either distinct or obscure; apex, fulvous amber; inside of aper- 
ture bluish-purple by reflected light, although no color is visi- 
ble by transmitted light. Whorls: 4% to 434; flatter and with 
sutures less deeply impressed than in P. straminea. Sculpture: 
weak, but with both series of oblique striae equally prominent ; 
although, in patches on the apical side, one set may dominate 
over the other, while, on the umbilical side, they cross each 
other so as to produce a malleate appearance. Aperture: oval; 
peristome simply and slightly thickened. Umbilicus: a little 
more than 4 the major diameter. Operculum: exterior, flat 
to slightly convex; interior with central boss less prominent 


than in P. straminea. 


The form from Palma Sola (fig. v-M) is considerably more 
elevated, shows more prominent corrugations, and is lighter in 
color than the typical form (figs. v-K, L). The apex is light 
amber, while the later whorls are brownish-amber above and 
amber below. ‘The bluish iridescence inside of the aperture is 


considerably lighter. 


From Pfeiffer’s original description, and Reeve’s figure 
(1865, Conch. Icon., vii-35b), which agrees in all particulars 


and apparently represents the type, P. g. glaucostoma is a very 


Occasional Papers of the Museum of Zoology 39 


similar shell to this subspecies. However, the typical form is 
considerably smaller and the color is more pronounced. 

I did not collect this form at Aroa, but a set in the A. N. 
S. P. (no. 12965), labeled as from that locality (C. F. Starke, 


1872), agrees quite closely with the shells from the Cerritos de 
Yumarito. 


Poteria dunkeri (Pir.) and approaching var. perezi (Hidalgo). 

: Figs. v-S, U. 

Four adults and 17 immature specimens; from the brook 
flats of a small tributary of Rio Lobaterita, near Estacién Ta- 
chira (H, I, b, 35), and from the steep valley-sides of the 
larger stream itself (H, I, c, 38). 

As will be seen from the table of measurements, these 
small lots of specimens show a great deal of variation, both in 
size and shape. All of the specimens are more elevated than 
either typical dunkeri or perezi. One specimen (fig. v-U) is 
much larger than the others, and, in this particular approaches 
P. d. dunkeri. The growth lines of all are well-marked, but 
regular and closely spaced. The peristome is thickend and often 
narrowly reflected, especially on the palatal wall. The number of 
whorls varies from 414 to 434; the umbilicus is from 1/3 to 
1/4 of the major diameter. The ground color varies from 
light olive-green to brownish-olive. The light, peripheral band 
is poorly marked, but is bordered below by a broad band of 
darker color, which tends to be accentuated at its upper and 
lower borders, or even to break up into a number of rather 
indefinite bands. In some specimens, an additional dark band 
is present on the umbilical side. as is characteristic of P. d. 
perezi, while the sutural region also may be darkened. 

Four specimens, labeled Venezuela (A. N. S. P. 12971), 


connect this series with typical P. d. dunkeri. Their measure- 


40 University of Michigan 


ments are given in the accompanying table. Besides the us- 
ually more depressed form, P. dunkeri differs from P. quiten- 
sis (Pfr.) in color (see key). In addition, the growth lines 
of the latter are developed into plications in the umbilical re- 
gion. However, P. gigantea (“Gray” Sowerby) has both red- 
dish and green color-forms, and P. gigantea fischeri (Hidalgo) 
shows the umbilical prominence of the growth lines. In real- 
ity, no very distinct characters, for the specific separation of 


many of the forms in the group of P. inca (Orbigny), have 


been found as yet. 


Measurements in the Group of P. inca 


Altitude Maj. Diam. 
P. d. dunkeri 17.8 198 (35.2) Conch. Cab. (1912) ; fig. 140-14. 
ASN Ss a0 78 
Means: (4specimens) 17.6 077, (3509 Venezuela. 
Extremes: ditte 15.7-19.9 170-195 (29.6-34.6) 
H, I, b, 35 (fig. v-U) 20.7 168 (34.8) Estacion Tachira, Ven. 
12.0 177 (213) 
P.d. perezt 13.6 180 (24.5) Figure, Hidalgo (1866). 
H,- Pabeese 14.3 17I (24.4) Baeza, Ecuador. 
(fig. v-S) 18.0 160 (28.8) Estacion Tachira, Ven. 
P.d.cardozi, H, 1, b, 40 133 159 (21.2) La Fria, Venezuela. 
10.8 182 (19.7) 
(type: fig. v-R) 9.4 181 (17.0) 
P. dunkeri var.; H,I,b,40 13.1 163)(21-3')) Quebrada Santa Aguita, Ven. 
14.7 159 (23.4) 
(fig. v-T ) 16.5 146 (24.1) 
P.p..popayana 15 136 (21.0) Figure, Lea (1838). 
A.N.S. P. 12980 17.8 122 (21.7) “New Granada.” 
A. N:S. P..12073 
Means: (3 specimens ) 13.4 146 (19.5) Cucuta, Colombia (Penny). 
Extremes : ditto 1I.7-14.9 139-152 (17.8-21.8) 
A, NiOAE. 12077 
Means: (7 specimens) 16.9 149 (25.1) Mendez, Colombia (T. Bland). 
Extremes : ditto 14.5-18.7 125-157 (22.6-27.1) 


P. p. fasciata(2 shells)16.5-17.0 141-158 (24.0-26.0) 


A. N. §. P. 12972 (12974) 
Means: (16 specimens) 17.0 


139 


(23.1) 


Extremes : ditto 14.6-18.7 121-153 (20.7-26.2) 


Kobelt (1922) Puerto Cabello. 


Puerto Cabello (C. F. Starke). 


Occasional Papers of the Museum of Zoology 4 


The following, very artificial key, to the names in the group 
of P. inca, is presented. The remarks, preceding the key to 
the group of P. translucida, also apply here. As the earlier 
descriptions are very monotonous reading, and often difficul: 
of interpretation, the characterization of each fresh lot as a 


new species appears to be the customary procedure in this 
group. 


A. Basal lip of peristome with a marked indentation, corresponding 
to subcanaliculate last whorl (teratological?; Cf. von Martens, 


1890). 
B. With a dark, spiral band just below periphery and another on 
base of the last whorl. Province Cauca, Colombia (about 


LS ele (OM i Ne POR 6 ol Ge) A Uae d= a 
Ey tahgliolatinn isl eiarston Paco Neocyclotus caucaensis DaCosta (1901.) 


BB. No bands mentioned in descriptions. 


C. Smaller, distinctly flattened, less vaulted. Bay of Salinas, 
Costa sled. 9 Tes ON SOAs, ton tebe nin bet naaee is als. owe 
rae Ee (!) Cyclotus irregularis, var. pittieri Mart. (1890). 


CC. Larger; the oldest name. Costa Rica. 26; 162(42); 5 (von 
Martens, 1890).....(*) Cyclostoma irregulare Pfr. (1855). 


AA. Basal lip of peristome normally entire. 
D. Periphery of last whorl angulate or subangulate. 


E. Obscure, cord-like development on periphery of last whorl; 
color olive-brown, somewhat lighter below. Ambato, 
Ecuador. 13.2; 163(21.6) ; 4%. Cyclotus past Crosse (1866). 


EE. Angulation more prominent; last whorl at least subcar- 
inate. 


F. Shell much depressed; growth-lines very prominent. 


G. Most depressed, rapidly increasing whorls; castaneo- 
fulvous with blackish carina. Colombia. 18.1; 202 
(36.6); 41%4...Cyclostoma laxatum Sowerby (1849). 


GG. Depressed; olivaceous with light, peripheral stripe and 
darker band below it. Bogota, Colombia. 20; 180(36). 
...(!) Cyclotus filoliratus Sowerby (1892, P. Z. 5.). 


FF. Less depressed shell; growth lines fine above, but coarse 
below (Cf. fischeri). 


42 University of Michigan 


H. Shell shaped like P. gigantea; blackish fuscous with 
vellowish brown, peripheral stripe. Ecuador. 23; 


BOOB 3S) FAY she ose Sia aod ope Se, unten axe e oie ee 
...N. (giganteus var?) subcingulatus Kobert (1912). 


HH. Shell more elevated; olivaceo-castaneous, with light 
stripe at periphery and darker band below. Colombia. 


20:5 = TAA CAS) A AUS. 0) acces eas eG oe Eee 
ah Set hs Seg Cyclostoma cingulatum Sowerby (1843). 


DD. Periphery of last whorl rounded. 
I. Shells very large; major diameter more than 45 mm. 


J. Last whorl descending, with upper angle canaliculate; 
dark band below periphery. Vera Paz, Guatemala (?) 


BU oT OAS 7 enh cites ous cpakidinis tis oo cee ae ae eee 
..(!) Aperostoma confusum Sykes (1901) Jour. Malac. 


JJ. More depressed; descent of last whorl less pronounced. 


K. Color rich chestnut-brown; lighter at suture and base, but no 
spiral bands. Zaragoza, Colombia (L 7, 75). 32; 159(51); 5. 


KK. Dark, spiral band below lighter stripe at periphery. 


L. Slightly more elevated; growth lines much coarser towards 
umbilicus than on sutural side of last whorl. Ecuador. 
82. IGECE HO) Ss Staaten Cyclotus fischeri Hidalgo (1867). 


LL. Most depressed; growth lines not markedly coarser belcw. 
Near Panama City. Panama: 363° 166(50:8))> 532.2 eee 
eae AS Cyclostoma giganteum “Gray” Sowerby (1843). 

(+ C. cumingti Jay, 1850). 


II. Shells smaller; major diameter less than 40 mm. 


M. Much depressed, rapidly increasing whorls; color brownish- 
olive, with light stripe above periphery, bordered by dark 
brown above and below. Peru. 13; 212(27.5); 4. ......-cses 
See Sete ee Neocyclotus depressus DaCosta (1906). 


MM. Less depressed; not more than twice as broad as high. 


N. Color predominantly reddish; usually distinctly larger and 
heavier shells, with prominent growth lines, at least 
around umbilicus. 


O. More depressed; chestnut-brown above, dark-brown be- 
low, with narrow, peripheral stripe of light color; 
growth lines weak. Pozuzo, Eastern Peru. 20.4; 183 
(38) 3) 434: <r Neocyclotus peruvianus DaCosta (1906). 


Occasional Papers of the Museum of Zoology 43 


OO. Less depressed, major index around 160. 


P. Rich chestnut in color with yellowish streaks and 
darker lines at irregular intervals, but slight trace of 
bands; growth lines prominent. Las Nubes, 4000 ft., 
Santa Marta Mts., Colombia (L 11, 74). 203; 157 
(CSC) ALG ee hic cadre eater ate sieges si (!) Aperxostoma 
sanctaemarthae Pilsbry and Clapp (1902) Nautilus. 


PP. Fulvous chestnut, with light, peripheral stripe, usually 
bordered below by dark band. 


Q. Slightly more depressed; black band above and 
below peripheral stripe; apex rose. Near Bogota, 
COOMA ee OK MOT (BBS isiuGa, yi - es = clea aie ale! acys ocsers 

Neocyclotus colombiensis DaCosta (1901). 


€.0) o> eins 


QQ. Slightly more elevated; greenish around umbili- 
cus; growth lines plicate below. Quito, Ecuador. 


BT Ot Tin ORAL) SAL amr pe vata cha belo sy ony Sai ore avis eels 
Cyclostoma quitensis Pfeiffer (1851). 


i 


NN. Color predominantly greenish; usually smaller and thinner shells 
(but see dunkeri and corpulentus). 


R. Much depressed shells; major diameter index over 180. 
S. Large and heavy shells. 


T. Last whorl swollen, almost solute; color dark olive 
brown with numerous, dark, spiral lines; growth lines 
prominent. San Sebastian, Colombia (LI, 11, 74). 10; 
184(35). (*) Cyclotus corpulentus E. A. Smith (1878). 


TT. Last whorl normal; color olive with light, peripheral 
stripe, bordered by dark band below. New Granada; 
Riobamba, Ecuador. 17.8; 108(35.2); 434. ......---: 
FOA.6 O06 bOI ae eRe Cyclotus dunkeri Pir. (1856). 


SS. Smaller and thinner shells; fusco-olivaceous with black 
band around periphery and another at base. 


i Lareer.. Baeza, Ecuador, Tae isdtes 5) 42 Foot. 2s 
OCG Cte OT Te Cyclotus peresi Hidalgo (1866). 
UU. Smallest. La Fria, Venezuela. 9.4; 181(17.0); 3%. 
more) hoa 5S COO GEOL REO (!) P. dunkeri cardozi new. 


RR. More elevated shells; major diameter less than 170(30) ; olive- 
brown with light, peripheral stripe usually bordered by dark 
band below. 


V. Major diameter index around 160. 


44 University of Michigan 


W. Shell malleate below; brownish green. Bogota, Colombia. 
14; 157(22); 4%. ...Cyclostoma bogotensis Pir. (1853). 


WW. Growth lines fine and quite regular; olive brown. 


X. Aperture normal; shell slightly more (or less) de- 
pressed: ‘Boltyvia. 1095-.163(31) ; 5)... 4-Seeeee 
Pein st Se ka ME Cyclostoma inca Orbigny (1833). 
Fa vise fre See (+ C. colombiensis “Fer.” Orb., 1835). 

(+ C. nobile “Fer.” Pot. and Mich., 1838). 


XX. Original figure (poorly drawn) shows somewhat so- 
lute aperture and slightly more elevated shell. Cox- 
oeira. Woods, Bahia; Brazil. 18:3; 155(28.3) -ceeeee 

Cyclostoma blanchetianum Moricand (1838). 


VV. Major diameter index varies around 140. 


Y. Bands more pronounced. Puerto Cabello, Venezuela. 


17:0; 137( 23:1) 34-5 (See taple). <t.<5 ooo eee 
N. popayana fasciata Kobelt-Schwanheim (1912). 


YY. Dark, spiral band. Popayan, Colombia. 15.5: 136(21) ; 
17 SaaS Cyclostoma popayana Lea (1838). 


YYY. Lwo teddish bands: 11.62°129(4!9)i5) 4-5... eee 
Salt ass Ai Cyclostoma inconspicuum Sowerby (1843). 


Poteria dunkeri cardozi, new subspecies 
Fig. v-R. 


Three adults and 10 immature specimens from the lowland 
forest around La Fria (H, Il, b, 40). These specimens from 
the rich, luxuriant jungles are, peculiarly enough, considera- 
bly smaller than the remainder. The description of the type 
(fig. v-R) of this form follows: 

Shell: depressed, thin, translucid. Growth lines: regular, 
compressed. Color: brownish horn, apex tinged with fulvous ; 
periphery of last whorl lighter, bordered below by a diffuse, 
brownish band, which is somewhat darker at its upper and 
lower borders; another, narrower band on the umbilical side. 
Whorls: 334; flattened above and rapidly increasing. Umbili- 


cus: a little more than ™% the major diameter. Aperture: 


Occasional Papers of the Museum of Zoology 45 


oblique, transversely elliptical. Peristome: thickened and 
slightly reflected, especially on the palatal wall. 

While this variety seems simply a dwarfed form of P. dun- 
keri, it appears to be at least an ecological race. While the 
type is clearly an adult shell, the smaller number of whorls 
appears to indicate arrested development as well as actual re- 
duction in size. The largest adult from this locality is consid- 
erably more elevated, and thus approaches the next form in 


shape. Both of the larger shells have 414 whorls. 


Attention is called to the fact that the type of this form 
is less than % as large as P. dunkeri, or the largest shell in 
the Estacion Tachira series; while these latter are almost 2/3 
as large as P. gigantea (Sowerby). The correspondence of 
P. dunkeri and P. quitensis with the forms of P. gigantea and 
P. q. fischeri, has already been noted. In addition, the last two 
forms usually have a larger number of whorls than do the 


smaller shells. 


Poteria dunkeri, var. approaching P. popayana (Lea). 
Fig. v-T. 


Three adults and 8 immature specimens from the creek flats 
of Quebrada Santa Aguita, near La Fria (H, I, b, 42). These 
shells are quite heavy and are considerably more elevated than 
most of the others from the same general region. The ground’ 
color of the adults is dark brown, so that the spiral banding 
is partially obscured. The shell substance at the apex is rosa- 
ceous to coral red. The aperture is nearly circular, and the 
peristome is thickened but not at all reflected. The umbilicus 
is about 4 the major diameter. 

As will be seen from the table of dimensions, this form ap- 


proaches P. popayana in shape. However, in the specimens. 


46. University of Michigan 


measured, the umbilicus of the latter is considerably smaller, 
around 1/5 the major diameter, although that of a very much 
elevated specimen from “New Granada” (A. N. S. P. 12950) 
is about 2/9. Von Martens (1873) has called attention to the 
intergradation in shape between P. inca and P. popayana. He 
has also pointed out that actual specimens of C. blanchetianum, 
obtained through Moricand, are more depressed than P. inca, 
instead of more elevated, as would appear from Moricand’s 
figure (see the preceding key). Specimens in the A. N. 8. P. 
(no. 12968, Brazil, from Moricand) agree with Von Martens’ 
statement. ‘To be perfectly candid, the difference in locality 
is about the only remaining excuse for the specific separation 
of the forms from the Brazilian region and those from the 


Colombian. 


Besides those already mentioned, measurements of sets in 
the A. N. S. P. from Cucuta, Colombia (L 8, 73), Mendez, 
Colombia (L 10, 75), and Puerto Cabello, Venezuela, are in- 
cluded in the table of dimensions. The extremes of variation 
in the major diameter index are especially noteworthy. Ko- 
belt-Schwanheim (1912), in the description of Poteria popay- 
ana fasciata, has called attention to the color variation in 
“Puerto Cabello” specimens, but apparently his series did not 
show as much variation in shape as the set in the Acad. of 
Nat. Sci. of Philadelphia. It is interesting, at least, that the 
larger forms, of the confusa-belli-fischeri-gigantea series, show 


similar differences in shape (see key). 
DIPLOM MATININAE 


Adclopoma occidentale (Guppy, 1872). 


Very abundant; at Quebrada La Fria, in rich leaf mould 


on an abandoned cacao plantation, that has returned (‘mon- 


Occasional Papers of the Museum of Zoology 47 


tado”) to practically the same conditions as the original forest 
(H, V, b, 41). Such isolated occurrence in second-growth 
seems to favor the hypothesis that this is an introduced species. 

The columellar denticle in these shells is quite low and 
rounded, but in no sense vestigial. The first 1% whorls are 
lightly and irregularly pitted. In addition to the prominent, 
growth ribs, the surface of their interspaces is covered with 
microscopic sculpture. The latter consists of rounded spiral 
riblets, which are broken, by impressed growth lines, into 


series of bead-like prominences. 


Figure 21. Adclopoma occidentale. -Teeth of half-row, separated 
laterad. At the right is shown a ventral view of another of the outer- 
most teeth. The scale indicates a length of 50 microns. 


Figure 21 shows a half row from the radula of this species, 
with the teeth separated laterad, so as to better show their 
form. As usually seen, the first lateral overlaps the central, 
while the bases of all of the laterals and marginals are crowd- 
ed closely together. The right-hand figure of the marginal 
tooth shows the ventral side uppermost. The denticle, near 
the center of the base of the rachidian tooth, is very promi- 


nent and distinctly reminiscent of the Amnicolidae. As _ will 


48 University of Michigan 


be seen from the scale (50 microns), some of the cusps ap- 
proach the limits of microscopic visibility. This radula ap- 
pears to be similar to that of Adelopoma tucma Doehring 
(1884); it certainly does not resemble those of the Apero- 


stomatinae. 


DESCRIPTION OF PLATES 


All drawings are made with the aid of the camera lucida. 
Under the number of each figure, is a hair line, which repre- 
sents an actual length of one millimeter in plate II, and 50 
microns (.05 mm.) in plates III and IV, and in the text-fig- 


ures. 


50 


University of Michigan 


PLATE I. 


All drawings were made with the aid of the camera lucida. The 


scales in figures I, 2, 3, 4, and 6 represent an actual length of I mm.; 
that of figure 5, Io microns (1.01 mm.). 


Figure 1. Leptinaria (Neosubulina) gloynii. 
Figure 2. L. gloynti. Apical whorls. 


Figure 3. L. gloynit. Last whorl, opened to show arrangement 
of columnellar and parietal lamellz. 


Figure 4. Microcramus bonairensis curacoana. Outline of 4 indi- 
viduals of type lot to show extreme variation in shape 
and size. 


Figure 5. M. b. curacoana. Radula; central and first laterals in 
place, 21st and 35th laterals. The curved line below 
the scale shows the shape of a single transverse row. 


Figure 6. Chondropoma (?) ravent. Apical whorls of young 
shell. 


A it 


Oar 21 


N 


» LO. 


University of Michigan 


PEATE UT 
The scales indicate a length of one millimeter. 
Lucidella venezuelensis. Front view of shell and outer view 


of operculum (calcareous plate). 


Tudora williamsoni. Outer view of operculum and front 
view of last whorl of type specimen (male). 


Tudora plicatula. Outer view of operculum. 


Tudora megacheilos. Outer view of operculum. 


VENEZUELAN SHELLS PEATE II 


54 University of Michigan 


PLATE At 


The scales indicate a length of fifty microns. The relationships of 
the centrals and laterals are represented as viewed. 


Fig. 11. Oligyra (Analcadia) rufa. Centrals, laterals, and tips of 
Ist, 4th, and roth uncini. 


Fig. 12. Oligyra (Analcadia) dysoni barbata. Centrals, and tips of 
Ist and 4th uncini. 


Fig. 13. Oligyra (Idesa) rotunda. Centrals, and top of Ist uncinus. 


Fig. 14. Oligyra (Sericea) riparia tachirensis, Centrals, laterals, and 
tips of Ist, 4th and 7th uncini. 


VENEZUELAN SHELLS PLATE III 


56 


The 


University of Michigan 


PLALE, 1V 


scales indicate a length of fifty microns. The relationships of 


ig. 18. 


. 19. 


the centrals and laterals are represented as viewed. 


Helicina (Oxyrhombus) concentrica. Centrals, laterals, and 
tip of 3rd uncinus. 


Helicina (Angulata) caracolla. Centrals and laterals. 


Lucidella (Lindsleya?) venezuelensis. Centrals, and tips of 
Ist and &th uncini. 


Helicina (Angulata) rhynchostoma ernesti. Centrals and tip 
of 3rd uncinus. 


Lucidella (Poeniella) plicatula. Centrals and tip of 3rd 
marginal. 


BS Ge ee ee 


University of Michigan 


PLATE V 
Reproduced approximately natural size. 


Poteria translucida translucida. la Guaira, Venezuela. 
Poteria translucida trinitensis. Male; Cariaquito, Venezuela. 
Poteria translucida trinitensis. Female; Cariaquito, Venezuela. 


Poteria translucida bejumensis. Type; female; station 7. 


Poteria translucida bejumensis. Most depressed female; station 7. 


Poteria translucida bejumensis. Smallest male; station 7. 
Poteria translucida bejumensis. Largest female; station 15. 
Poteria translucida major. Type; Caracas, Venezuela. - 
Poteria transiucida santaguitensis. Male; station 42. 
Poteria translucida santaguitensis. Type; female; station 42. 
Poteria glaucostoma aulari. Male; station 28. 

Poteria glaucostoma aulari. Type; female; station 28. 
Poteria glaucostoma aulari. Female; station 20. 

Poteria straminea. Female; station 3. 

Poteria straminea. Male; station 3. 

Poteria granadensis rugata. Female; Cariaquito, Venezuela. 
Poteria granadensis rugata. Male; Cariaquito, Venezuela. 
Poteria dunkeri cardozi. Type; male; station 4o. 

Poteria dunkeri, near peresi. Station 38. 

Poterta dunkeri, approaching P. popayana. Station 42. 


Poteria dunkeri dunkert. Largest female; station 35. 


VENEZUELAN SHELLS PLATE V 


NUMBER 138 JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


THE HABITAT OF APHREDODERUS SAYANUS IN 
KALAMAZOO COUNTY, MICHIGAN 


By Hersert Ray BECKER 


The first published note of the occurrence of the pirate 
perch (Aphredoderus sayanus) in Michigan was made by 
Bollman,! who recorded “two specimens taken in a small 
brook which flows into Howard Lake.” Although the several 
small streams (only a few rods in length) which are tributary 
to this lake have often been examined by the writer, they 
have never yielded Aphredoderus. Eucalia tconstans, in 
contrast, lives here in great abundance. 

Aphredoderus sayanus is, however, a common fish in two 
streams flowing from the north into Black Lake, immediately 
north of Howard Lake. The larger (more easterly) of these 
two brooks rises little more than two miles from the lake in 
a level oak country. Although but a small ditch for most of 


its length, in its last third of a mile it expands into a gentle 


* Bull. U. S. Fish Comm., 8, 1888 (1890), p. 223. 


2 University of Michigan 


meandering stream six to fifteen feet in width, seldom more 
than three feet deep except in high water, and with a current 
flowing at the rate of twenty-five feet per minute. In mid- 
stream, the water here is practically free from vegetation, but 
for a quartér of a mile from the lake both banks show a 
profuse growth of water cress, together with small patches of 
peppermint (Impatiens biflora), Scutellaria lateriflora, Iris 
versicolor and Mimulus jamesti. Close to the lake the banks 
become impaszably boggy, with growths of cat-tail (7ypha 
latifolia), swamp loosestrife (Decodon verticillata), Spargan- 
ium and swamp dock (Rumesx brittanica). The bottom of 


the stream remains firm and sandy to its entrance into the 
lake. 


Aphredoderus is not found in the upper portion of this 
brook, where the water dashes over gravel—the habitat of 
the rainbow darter (Poecilichthys coeruleus). It is abundant 
however, in the lower quieter course of the stream. Here 
it is associated with an abundance of aquatic insects, amphi- 
pods and snails, and with twenty-four other species of fishes. 
These fishes are: Amia calva, Catostomus commersonni, 
Erimyzon succeta, Pimephales notatus, Semotilus atromacu- 
latus, Notemigonus crysoleucus, Notropis cornutus, Nocomis 
kentuckiensis, Ameiurus natalis, Ameiurus melas, Umbra limi, 
Esox vermiculatus, Fundulus notatus, Eucalia inconstans, 
Labidesthes sicculus, Ambloplites rupestris, Chaenobryttus 
gulosus, Lepomis incisor, Eupomotis gibbosus, Micropterus 
salmoides, Boleosoma nigrum, Poecilichthys iowae, Poeci- 


lichthys coeruleus, Microperca punctulata. 


The smaller tributary of Black Lake is three or four feet 
deep in its lower course. It has here a miry bottom covered 


in many places with a growth of Chara and Elodea, 


Occasional Papers of the Museum of Zoology 3 


and large patches of water cress occur on both banks. The 
liverwort Riccia fluitans forms a large part of the floating veg- 
etation while Marchantia is found along the shore. Here mol- 
lusks are more plentiful than in the preceding stream ; the same 
profusion of water bugs occurs here, and dragon fly nymphs 


are even more numerous. 


Aphredoderus is abundant in this lower portion of the 
stream, living in the cress in company with Umbra limi, 
Eupomotis gibbosus, Ameiurus, sp., Micropterus sa’motdes, 
and Eucalia inconstans. It is more loosely associated here 
with Notropis cornutus, Notemigonus crysoleucas and 
Pimephales notatus, these species holding to the open water 
in midstream. ‘Trout have been repeatedly planted in this 
brook, but none has been found during these investigations. 

Farther upstream, where the surface becomes almost 
covered by cress and peppermint, Aphredoderus was not 
found. 


Not all streams in the immediate vicinity of Black Lake 
contain pirate perches. In a large deep ditch near Updegrove 
Lake Aphredoderus is rare, but Eupomotis gibbosus, Microp- 
terus salmoides, Umbra lint, and Fundulus notatus are 
abundant. A stream flowing into Red Lake is too sluggish 
for Aphredoderus ; Umbra limi swarms beneath its duckweed 
covered surface. A brook tributary to a stream flowing into 
Barton Lake abounds with trout (Salvelinus fonttnalis), 
Umbra limi and Esox vermiculatus, but Aphredoderus is 
absent. 

Pirate perches are lowland fishes, in the region studied 
never being found far from large streams or lakes. They 
inhabit dense vegetation and avoid open water: over ninety 


per cent of the many specimens taken in this locality were 


4 Umversity of Michigan 


found beneath water cress or in masses of Chara. After the 
cress has been killed by the cold of winter, they still lurk in 
deep holes underneath the bank in company with Umbra lim, 
Eupomotis gibbosus, etc. They are distinctively bottom fishes. 

The food of the pirate perch was found to consist of 
certain of the smaller invertebrates of its vegetation habitat, 
namely green midge larvae, small amphipods and sometimes 
the larvae of certain dytiscid beetles. Many stomachs have 
been found empty. 

Pirate perches pass a considerable part of the time at rest: 
head upstream, the large dorsal expanded, the caudal partly 
closed and the pectorals slowly moving. When frightened 
they move swiftly ; liberated in midstream they dive vertically 


downward to find a place of concealment. 


NUMBER 139 JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arzor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NOTE ON THE GENERA COLUBER AND MAS''- 
COPHIS, AND A DESCRIPTION OF A NEW 
SPECIES OF MASTICOPHIS 
By Artuur I. ORTENBURGER 

A study of the new world snakes, commonly called racers or 
whip snakes, has made it evident that the American species of 
the genus recently known as Coluber (L.) form two natural 
groups sufficiently distinct to be called genera, as will be 
shown in detail in a forthcoming monograph. Masticophis 
and Coluber are the names to be applied to these genera. 
Since the new form described in this paper must be included 
in the genus Masticophis it is advisable to point out the prin- 
cipal characters of the two genera, which involve differences 
in the dentition, in the method of reducing the scale rows, 


and in the structure of the penis. 
Genus MastTIcopuHIs 


Diagnosis: Maxillary teeth solid, 16-23 in number, longer 


and stouter posteriorly, diastema present or absent; dentary 


2 University of Michigan 


teeth in general longer anteriorly, scaphiodont, but occasion- 
ally the last few smaller, hence sometimes kumatodont; head 
distinct ; cephalic plates normal; eye large, pupil round; scales 
smooth, in 17 or 15 rows, when 17, reduced to 13 or 12 by 
the loss of both lateral and dorsal scale rows, when 15, usually 
reduced to 12 or 11 by the same method, if reduced only to 
13, fifth lateral row lost; anal plate divided; form elongate; 
tail very long; caudals in two series; penis bilobed, sulcus 
simple, spines 30-75, and in 2-5 longitudinal rows, calyces in 


Q-14 rows. 
Genus CoLUBER 


Diagnosis: Maxillary teeth solid, 13-16 in number, increas- 
ing in size posteriorly, diastema usually present; dentary teeth 
strong and stout, subequal in general, longer anteriorly, but 
last few at each end of series smaller; head distinct; cephalic 
plates normal; eye large, pupil round; scales smooth, in 17 or 
I5 rows, when 17, always reduced to 15 by the loss of a lat- 
eral row on each side of the body, when 15, no reduction; 
anal plate divided; form elongate; caudals in two series; penis 
bilobed, sulcus simple, spines 90-135, and in 6-9 longitudinal 
rows, calyces in 6-8 rows. 


AMERICAN SPECIES OF CoLUBER AND MAS‘TICOPHIS 


Masticophis (B. & G.) ruthveni Ortenburger 
anthonyi (Stej.) schotti B. & G. 
aurigulus (Cope) semilineatus (Cope) 
barbouri (Van Denburgh) taeniatus girardi (Stej. & Barb.) 
flagellum flagellum (Shaw) taeniatus taeniatus ( Hallow.) 
flagellum flavigularis (Hallow.) 
flagellum frenatus (Stej.) Coluber (L,.) 
lateralis (Hallow.) constrictor constrictor (L,.) 
lineatus (Bocourt) constrictor flaviventris (Say) 
mentovarius (Dumeril & Bibron) constrictor mormon (B. & G.) 


piceus (Cope) oaxaca (Jan) 


——— se tal 


Occasional Papers of the Museum of Zoology a 


Masticophis ruthveni, new species 


Diagnosis: Characters very similar to those of Masticophis 
schotti (B. & G.) from which it differs primarily in possessing 


no dark lateral stripes. 


Range: Known only from Tamaulipas, Mexico, and ex- 


treme southeastern Texas. 


; Type Specimen: Cat. no. 57681, Museum of Zoology, Uni- 


versity of Michigan. Brownsville, Texas. 


Description of Type Specimen: Scales in 15 rows anterior- 
ly, reduced to 13, then to 12, and finally on the posterior por- 
tion of the body to 11; reductions as follows: 15-13 by loss 
of fifth row on each side of the body between scutes ITI-113; 
13-12 by loss of dorsal row above scute 131; 12-11 by loss 
of dorsal row above scute 142. Ventrals 193, anal divided; 
caudals 138 pairs; supralabials 8, the fifth extending up be- 
hind the eye; infralabials 9; oculars 2-2; temporals, right, 
2+ 3+ 4, left, 2 + 3 + 3; anterior chin shields in contact, 
shorter and wider than posterior, the latter separated by 4 
small scales; loreal long and narrow; nasals 2, nearly equal in 
size, the nostrils between them; cephalic plates very similar 
to those of M. schotti. 


Body slender and elongate, total length 168 cms.; tail 
length 52 cms. ; proportionate tail length 0.310; sex male; head 


long and narrow, length 42 mm., width 19 mm., proportionate 
width 0.45. 


The color and pattern are included in the general descrip- 


tion of the form, which follows. 


Remarks: This new species is very rare in collections, but 


it has been possible to study the coloration in a good series of 


4 University of Michigan 


living specimens. In the following description the nomencla- 


ture is capitalized when it is that of Ridgway. 


The general dorsal color is olive gray anteriorly, gradually 
changing to a reddish brown posteriorly. The actual variation 
in the series is from Olivaceous Black to Grayish Olive. The 
sides of the body vary from Deep Slate Olive to Grayish Olive. 
A distinct and very constant character of the species is the: 
yellow anterior edges of the dorsal scales; this is very evident 
if the skin is slightly stretched (Pl. 1). These yellow marks 
do not extend more than three or four scaie rows from the 


mid-dorsal scale row. 


The general color below is a bright yellow, anteriorly vary- 
ing from light Chalcedony Yellow to Shaw Yellow. This 
yellow changes to a light Blue Gray on the middle third of 
the body; on the posterior third, to pinkish; and, finally, near 
the tail in most specimens to a bright true Oid Rose—rarely 


to a Light Coral Red. 


The throat below is dotted with dark orange; the belly has 
very numerous minute gray dots. There is a concentration 
of the gray dots on the ventral surface toward the lateral one- 
fifth or fourth of the scutes, making this portion of the scutes 
a general gray color. The tail is immaculate. On some few 
specimens there is also a light yellowish lateral stripe occupy- 
ing the angle of the scutes and a small portion of row 1; 
the gray dotting may also break up so as to leave an un- 


dotted yellow stripe in a mid-ventral position on the scutes. 


’ 


The sides of the neck may have one or more light cream 
stripes. In one specimen, not fully grown, (M. C. Z. 13951) a 
light lateral stripe is present and continues the length of the 


body. Another specimen (U. S. N. M. 1974), very young,. 


Occasional Papers of the Museum of Zoology 5 


shows this light lateral stripe also, and except for the fact that 
the typical yellow marks on the dorsal scales are lacking it is 
quite similar to schotti. A few specimens after some years’ 
preservation in alcohol show faint lateral stripes anteriorly in 
positions comparable to those of schotti and other species of 
the taeniatus group; none show evidence of the distinct dark 
stripes found in schotti. 

The lateral scales, from a point just posterior to the angle 
of the mouth to about scute 10, are blotched with brilliant 
Brazil Red. The top of the head is Dark Grayish Olive to 
Grayish Olive; labials and chin shields pearl white to Chalce- 
dony Yellow; supralabials cream with dark upper edge of the 
same color as top of head, sometimes with a narrow lower 
border of gray blue; the iris is red-gold to gold-brown, 
always with a bright gold inner edge; upper and sometimes 
lower preocular with Straw Yellow spot; posterior nasal with 
a bright cream spot on anterior edge; chin shields usually 
faintly blotched or spotted with dark gray along sides. 

The dorsal scale formula is 15-13, 15-13-12, Or I5-13-I2-IT. 
Occasionally the last dorsal scale row lost is regained by the 
splitting of the row resulting from the fusion of the two dorsal 
rows just anterior. In such cases, the scale formula becomes 
15-13-12-11-12. The first reduction is always the result of 
the loss of lateral rows, the fifth at points about opposite on 
the body. The other reductions are due to the loss of one or 
more mid-dorsal scale rows. ‘The place of lateral reduction 
averages I17 scutes posterior; the first decrease in the dorsal 
series takes place 148 scutes posterior, the second, 161 scutes 
posterior. 

The ventrals vary from 188-197, average male, 193.4, aver- 


age female 194.2; the caudals from 131-149, (extremes, male 


6 University of Michigan 


131-149, female, 131-137), average male, 141.9, average fe- 
male, 134. The proportionate tail-lengths are, male, 0.326, 
female, 0.290. The supralabials are 8, the fourth and fifth 
entering orbit, the fifth being prolonged up and behind the 
eye. The infralabials are usually 9, often 10; the oculars 
2-2; the temporals usually 2 + 2+ 2 (Pl. I, Fig. 1), except 
that there are often 3 in either the first, second, or third series, 
and sometimes 4 in the second or third series. The posterior 
chin shields are longer than the anterior, the former being sep- 
arated by 3-5 small scales; the loreal is longer than high. 
(Fi-d1, Fig.2;) 


The head is long and narrow, the width (just posterior to 
the eye) divided by the length averages 0.43. 


The largest specimen examined is 168 cms. in total length, 


of which the tail comprises 52 cms. 


The penis exhibits the following structure: Strongly bifur- 
cate, the length of the forks being about one-seventh the total 
length of the organ; sulcus simple, extending to the tip of the 
penis on the side of the larger lobe; distal surface smooth, 
surrounded by rather deep, strongly-fringed calyces, which 
extend proximally in about 9-10 rows, calyces covering between 
a third and a half of the entire organ; the fringe of the calyces 
at a point about halfway to the base, rapidly changing to dis- 
tinct, small spines, which become larger toward the base of 
the organ; spines 43-46, in 3-4 (sometimes 5) rows; three 
large basal spines proximad of general spined portion; region 
along the sulcus, through the spined area, thickly covered with 
spinules (Pl. III). 


The dentition, as shown by several specimens, is as follows: 


maxillary teeth 16-17, coryphodont, dicranterian, the last 


Occasional Papers of the Museum of Zoology q 


three teeth separated by a diastema; dentary teeth 21-23 in 
number, usually 21, scaphiodont, uniformly increasing in size 
anteriorly, the first few being nearly twice as long as the last; 
palatines 13-16, usually 13 or 14, only slightly longer anterior- 


ly; pterygoids 18-25, usually 22-23, scaphiodont. 


It should be pointed out here that it is entirely possible that 
this new form, here given the status of a species, may later be 
found to be a subspecies of schotti. It is impossible to deter- 
mine this, however, until specimens from critical localities are 


available. 


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Masticophis ruthveni Ortenburger. Cat. No. 57687, 
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MASTICOPHIS RUTHVENI PLATE JII 


NUMBER 140 JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arpor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NEW NORTH AMERICAN SNAKE, OF THE 
GENUS NATRIX? 


By Frank N. BLANCHARD 


No systematic treatment of the North American species of 
Natrix akin to the Coluber fasciatus of Linnaeus has been sat- 
isfactory, and the present reviewer is under no illusions as to 
the permanence of his present views on the subject. It is 
time, however, to point out the unity and relatively restricted 
range of fasciata and its nearest relatives, and the marked dis- 
tinction between the southeastern and the Mississippian phases. 
For the latter there is apparently no name available, and it is 


therefore proposed that it be called: 


Natrix fasciata confluens, new subspecies 


1892 Tropidonotus obliquus Garman, S., Bull. Essex Inst., vol. 24, 
p. 6. 


*Contributions from the Department of Zodlogy, University of 
Michigan. 


2 University of Michigan 


1908 Tropidonotus sipedon fasciatus Strecker, Proc. Biol. Soc., Wash- 
" ington, vol. 21, p. 50; same, pp. 69, 76; Baylor Bull., vol. 18, 
no. 4, 1915, p. 28. 


1911 Natrix fasciata Hurter, Trans. Acad. Sci., St. Louis, vol. 20, 
no. 5. Pp. 154. 
Diagnosis: Similar in scutellation and proportions to Na- 
trix fasciata fasciata (Linnaeus), but with the dorsal saddles 
very much larger and only about half as numerous, and with 


average differences as shown in the table, page 5. 


Range: Eastern Louisiana north through southern and east- 
ern Arkansas to southeastern Missouri, and west in Texas to 


about the 98th meridian (See map, page 7). 


Type specimen: Museum of Zoology, University of Mich- 
igan, number 57707; Butler County, Missouri, April 16, 1905; 


3, Thutter, collector, 


Description of type specimen: Ventral plates, 135; anal plate 
divided; caudal plates 70, all divided; dorsal scale rows, 21 on 
the anterior portion of the body, changing to 23 (on the left 
side at the level of the 30th ventral plate, by addition, appar- 
ently, of a 5th row, and on the right side at the level of the 
33rd ventral scute, by addition, apparently, of a 6th row), 
then to 21 (on the left side at the level of the 67th ventral, 
and on the right side at the level of the 63rd ventral, by loss 
of the 5th row, apparently in each case), then to 19, and to- 
wards the posterior end to 17, the full formula being, there- 
fore, 21-23-21-19-17; upper labials 8 on each side; lower 
labials, 10 on each side; 1 preocular; 3 postoculars, the lowest 
of which is on each side much the smallest; a single anterior 
temporal on each side, followed by 3 posterior temporals ; pos- 
terior chin shields a little longer than the anterior, separated 


anteriorly by about the width of one small scale and diverging 


Occasional Papers of the Museum of Zoology 3 


posteriorly ; other head shields normal for the genus; all of the 
dorsal scales keeled, but the first row less strongly than the 
others. 

Total length, 558 millimeters; tail length, 130 millimeters; 
tail, therefore, 0.233 of total length. Sex, male. 

The general color above is a very dark brown. Crossing 
this, more or less transversely, are about 13 light yellowish 
bands, one scale wide. ‘These are more or less mottled with 
darker, especially posteriorly, where, also, they are mostly in- 
terrupted on the median dorsal line. The belly is rather heav- 
ily checked with black, especially posteriorly, and the under 
side of the tail is almost uniformly black. The head is dark 
brown above, light brown on the sides, and immaculate light 
yellow beneath. The labials are almost entirely unmarked, and 
the chin shields, gulars, and anterior ventrals are quite so. On 
the postoculars, temporals, and last two upper labials are a 
few mottlings representing the lower border of the light post- 


ocular band of N. fasciata fasciata. 


Remarks: ‘Thirty-one specimens have been examined, rep- 
resenting the following localities: Butler and Dunklin coun- 
ties, Missouri; Miller and Jefferson counties, and Wheatley, 
Arkansas; Jefferson County, Mississippi; Jackson County, and 
New Orleans, Belair, Prairie Mer Rouge, and Avery Island, 
Louisiana; Brazoria, Cook, and Falls counties, Dallas, and 
Angelina River, Texas; and “New Orleans to Galveston.” 
Published records that appear to refer definitely to this form 
name the following additional localities: Victoria, Tehuacana 
Bottoms, Laguna Lake, and Demings Bridge (Matagorda 
County), Texas; Hot Springs and Texarkana, Arkansas; and 
Stoddard County, Missouri, (See references in the 


synonomy. ) 


4 University of Michigan 


These specimens indicate a well-marked race, always recog- 
nizable at a glance by the peculiar pattern. The latter seems 
to have resulted from a fusion by twos of the dorsal blotches 
or saddles of fasciata. Furthermore the postocular light band 
is very prominent, and its lower dark border may be reduced 
to practical absence. Specimens from southeastern Louisiana 
show the closest relationship with fasciata. Indeed a few 
specimens examined by the writer and labelled “New Orleans” 
must be identified as N. fasciata fasciata, and Mr. Percy Vios- 
ca informs me that in this region both phases occur in the 
same localities in “almost infinite variation, sometimes appar- 
ently in the same brood.” New Orleans is therefore within 


the region of subspecific intergradation. 


Other characteristics of this form are tabulated below. 


Discrimination of the subspecies of the fasciata group :— 
Perhaps the most constant feature by which the subspecies, 
N. fasciata fasciata, N. fasciata confluens, and N. fasciata pic- 
tiventris may be distinguished from other species of Natrix 
with similar scutellation is the light yellowish or brownish 
band extending backwards from the eye to the angle of the 
mouth. A specimen is rarely so melanistic that wetting the 
head will not reveal traces of this feature, and specimens of 
confluens may have it obscured only by its broadening and the 
practical disappearance of its lower dark border. But within 
the fasciata group it is not always so easy to distinguish N. 
fasciata fasciata from N. fasciata pictiventris. Indeed there 
is a real temptation to synonomize the latter with the former. 
Numerous average differences appear, however, which lead the 
writer to believe that more thorough study on large series of 


specimens will amply justify the separation here maintained. 


Occasional Papers of the Museum of Zoology 5 


For purposes of comparison of these three races a table of 
extremes and averages has been prepared for certain of the 
features which show a significant degree of subspecific con- 
stancy. 


Summary of certain characteristics of the fasciata group 


Ventrals : 
Subspecies Males Females 
no. extremes average no. extremes average 
confluens 13* 129-138 134.0 18 128-138 134.1 
fasciata 22 126-137 129.6 26 127-133 130.6 
pictiventris 21 123-129 125.7 32 121-131 126.2 
Caudals 
Subspecies Males Females 
no. extremes average no. extremes average 
confluens 13 67-81 74.9 13 63-67 Gyes 
fasciata 15 70-83 Tea 24 63-76 68.8 
pictiventris 13 77-89 82.4 17 65-78 71.9 
Tail length divided by total length 
Subspecies Males Females 
no. extremes average _ no. extremes average 
confluens 13 .226-.265 .248 13 .212-.242 22, 
fasciata 14 .234-.291 L277 22 .193-.270 .242 
pictiventris 13 .260-.298 .279 17 .229-.281 -253 


Maximum number 
of scale rows 


Dorsal blotches on body Males Females 
no. extremes average 25 23 ae 23 
confluens 29 10-19 13.4 I 12 2 16 
fasciata 41 19-33 23.6 2 20 5 15 
pictiventris 40 24-35 29.1 4 17 23 9 


Fasciata, it will be seen, is strictly intermediate between con- 
fluens and pictiventris, except that in proportional tail length 
it is much closer to pictiventris. In general appearance also 
these two are much more like each other than either is like 
confluens. 


* Number of specimens examined. 


6 University of Michigan 


The more useful features for the separation of these races 


are embodied in the following synopsis of the fasciata group: 


a, Dorsal saddles on*pody about 11 to 9722. +2-<2-+4-% -..6-> ae 


RP Se Sethe SA ee PS ope Natrix fasciata confluens Blanchard. 
(Eastern Louisiana north to southeastern Missouri, eastern and 


southern Arkansas, and west in Texas to about the 98th meri- 
dian). 


a, Dorsal saddles on body about 20 to 33. 


b, Dorsal saddles on body commonly about 24; ventral plates us- 
ually more than 128; belly often with dark quadrate spots; 
often small lateral spots alternating with the dorsal saddles. 
Li tintle ste ncae oe ces ee dees Gia afascigta ‘fascia. tones 

(Northern Florida and the coastal regions from North Carolina 
to southeastern Louisiana.) 


b, Dorsal saddles on body commonly about 29; ventrals usually 
“less than 128; belly with dark, sometimes reddish, anterior 
borders on the ventral scales; often reddish markings with 
black edges particularly on the ends of the ventrals; no small 
lateral alternating spots.....Natrix fasciata pictiventris Cope. 
(Peninsular Florida.) 


ACKNOWLEDGMENTS 


The writer’s appreciation for loans of specimens used in 
this study, and for other courtesies, is cordially extended to 
Professor A. G. Ruthven of the Museum of Zoology of the 
University of Michigan, to Dr. Leonhard Stejneger of the 
United States National Museum, to Dr. Thomas Barbour of 
the Museum of Comparative Zoology, to Dr. John Van Den- 
burgh of the California Academy of Sciences, to Mr. KE. Ds 
Bunker of the Kansas University Museum, to Mr. H. P. Lod- 
ing of Mobile, Alabama, and to Mr. Percy Viosca, Jr., of New 


Orleans. 


*(SO[IIID ASART) SIN, {UOJ DyDIISD{ “AJ PUB *(So[DATD {[eUs ) 
stJuaeyIid VjolIsp, “No *(saienbs) vyvlosp{ vivlosv{ LiAjDAY JO} PsOdIL ApLoo] Surmoys duyy 


J My 


Sy 


pulyerely, 


NUMBER I4I JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arsor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NEW SPECIES OF PHYLLOPOD 
By G. S. Dopps 


Branchinecta occidentalis, new species 


Description: Male-—Length 9 mm. Swimming appendages 
eleven pairs; caudal segments without appendages, nine; cau- 
dal stylets a little larger than the combined length of the last 
two segments. Claspers (2nd antennae) a little longer than 
the distance between the outside surface of eyes (Fig. 1a); 
first and second segments of about equal length; first segment 
stout, its length about twice its thickness, moderately bowed, 
with concavity inward; this segment bears on its anterior, me- 
sial aspect, near the base, a good sized process, curved like a 
blunt horn, and bearing on its mesial (convex) surface a num- 
ber of spines; the two processes are placed with their apices 
close together, so that the two together make a prominent point 
between the bases of the appendages. Second segment much 
more slender than the first, slightly curved and gently taper- 


ing; the inner (concave) surface bears numerous small corru- 


2 University of Michigan 


gations. An abrupt offset on outer margin marks the joint 
between the first and second segments. 

The two penes (Fig. tb) are straight, slightly divergent, 
square in cross section, and extend about to end of second 
caudal segment. Each bears, on its mesial margin, a stout, 
curved spine, bearing three or four small teeth on its convex 
(mesial) margin. ‘These two spines are so placed that a near- 
ly circular area is enclosed between them and the bases of the 


two penes. 


Female—Length 10 mm. First segment of second antenna 
about as long as in the male, but more slender, its length be- 
ing a little more than three times its thickness (Fig. 1c). The 
segment is nearly straight and tapered somewhat in distal 
third; it bears on its mesial surface, near the middle, a small 
thumb-like process extending parallel to the appendage. The 
second segment is represented by a slightly curved spine, which 
arises from the inner, distal angle of the first segment. This 
spine and the thumb-like appendage of the first segment are 
of about the same size and proportions. 

The ovisac is attached to the ventral surface of the elongated 
second caudal segment, and has the form of a compact sac, 
extending very little beyond the caudal end of the segment. 
In 22 specimens examined, it never contained more than six 
eggs. 

In other respects the female closely resembles the male. 


Type Specimens: Cat. No. 52030, Museum of Zoology, 
University of Michigan; Collected on Stanford Campus, Palo 
Alto, California, by Professor Harold Heath, April 10, 1922 


mame 


Occasional Papers of the Museum of Zoology 3 


Fig. 1. FPranchinecta occidentalis: a, front view of head of male; 


b, ventral view of penes of male; c, front view of head of female; 


d, side view of Ist, 2nd and 3rd caudul segments and ovisac of female. 


' cae = jo! 

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NUMBER 142 JULY 9, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arzsor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


COMMENTS ON RING-NECK SNAKES (GENUS 
DIADOPHIS), WITH DIAGNOSES OF 
NEW FORMS? 


By Frank N. BLANCHARD 


In the course of a study of the ring-neck snakes, genus 
Diadophis, it has been found necessary to alter considerably 
the present systematic arrangement of the western forms and 
to propose several new names for the designation of subspecies 


hitherto unrecognized. 


Diadophis regalis regalis (Baird and Girard) 
1853 Diadophis regalis Baird and Girard, Cat. N. Amer. Rept., pt. 1, 
Serpents, p. II5. 

A study of all the specimens and records of ring-neck snakes 
from western Texas to Arizona makes it appear highly prob- 
able that two races inhabit this territory, an eastern form, es- 
sentially without the neck ring, ranging to southeastern Ari- 


1 Contribution from the Department of Zoology of the University 
of Michigan. 


2 University of Michigan 


zona, northern Sonora, and probably southern Utah. That the 
case is not wholly clear is readily admitted for there is some 
evidence that ringed specimens may occur in the eastern range, 
but the geographical distribution of the ringed and ringless 
specimens is so suggestive of two races that it is deemed ad- 
visable to open the question by the proposal of a name to rep- 
resent a western form of D. regalis distinguishable from an 
eastern form by the presence of a neck ring. This western 
race may be known as 


~ » 
Diadophis regalis arizonae, new subspecies 


Diagnosis: Like Diadophis regalis regalis Baird and Girard 


except for the possession of a broad ring of lighter color be-. 
hind the head. 


Type Specimen: United States National Museum, number 
625068, collected by G. Hofer. 


Type Locality: Sabino Cafion, Arizona. 
Range: Central Arizona to northern Sonora. 


Description of Type Specimen: ventral plates, 219; anal 
plate divided; caudal scutes, 65 pairs; upper labials, 7; lower 
labials, 8; preoculars, 2; postoculars, 2; temporals, I+-I+2 
(labials, oculars, and temporals the same on each side) ; dor- 
sal scale rows, 17, changing near the posterior end of the 
body to 15. General color above (in alcohol) a deep glaucous- 
gray,” extending on the sides over about half of the lowermost 
row of scales; below cream color forward over the chin and 
including the lower portions of the upper labials, suffused with 


reddish towards the posterior end of the body and under the 


2Colors are based on Ridgway’s Color Standards and Color No- 
menclature, I912z. 


Occasional Papers of the Museum of Zoology 3 


tail; neck ring 2% to 3 scales wide, ivory yellow, margined 
before and behind with black; head above much darker than 
the general dorsal surface; lower labials, chin shields, and ven- 
tral scutes prominently spotted with black; caudal plates with 
black extensions from their postero-lateral corners. Total 
length, 591 millimeters; tail length, 109 millimeters. Sex, 


male. 


Diadophis amabilis amabilis (Baird and Girard) 


1853 Diadophis amabilis Baird and Girard, Cat. N. Amer. Rept., pt. 1, 
Serpents, p. 113. 


The specimens from San Jose, California, to which the name 
D. amabilis was first given were originally regarded as repre- 
senting a form distinct from the only other then known speci- 
men of the genus from this state, but later writers have more 
often regarded these two as the same, and, indeed, have in- 
cluded under this name all specimens subsequently found in 
the west coast states. That this was an assemblage of several 
distinct but related races was early evident to the writer, and 
now, after a study of all the material available, it is evident 
that not only must the two forms originally described be re- 
garded as distinct but that, in addition to these, four others 
must be recognized if we are to attempt an understanding of 
the genus in this region. 

The subdivisions of this complex are based largely upon the 
number of dorsal scale rows, width of neck ring, maculation 
of the ventral surface, and the extent of the encroachment of 
the ventral color upon the dorsal scale rows. 

The name D. amabilis amabilis is here restricted to those 
snakes of the genus inhabiting the San Francisco Bay region 
and the valleys of the San Joaquin and Sacramento rivers. 


‘The characteristics of this race are a narrow neck ring that is. 


4 University of Michigan 


sometimes interrupted, dorsal scales in 15 rows often becom- 
ing 13 towards the posterior end of the body, ventral color 
usually extending onto the lower part of the second row of 


dorsal scales, and numerous small black spots scattered over 
the belly. 


Diadophis amabilis similis, new subspecies 


Diagnosis: Similar to D. amabilis amabilis, but with the 
light color of the ventral surface extending over only one-third 
to two-thirds of the lowermost row of dorsal scales. Cther 
distinctive features are the moderate amount of black spotting 
on the belly, the dorsal scales in 15 rows throughout, or drop- 
ping to 13 towards the posterior end of the body, and the gen- 


erally light olive color of the dorsal surface. 


Type Specimen: Museum of Zoology, University of Mich- 
igan, number 57897, collected by L. M. Klauber in the spring 


of 1923. 
Type Locality: Carmel, Monterey County, California. 


Range: Yrom southwestern San Bernardino County south 


into the San Pedro Martir Mountains in Lower California. 


Description of Type Specimen: Ventral plates, 189; anal 
plate divided; caudal scutes in 59 pairs; upper labials, 7; low- 
er labials, 7; preoculars, 2; postoculars, 2; temporals, 1-+1--2 
(labials, oculars, and temporals the same on each side) ; dorsal 
scales in 15 rows, becoming 13 near the posterior end of the 
body; neck ring 1% scales wide, not interrupted; ventral color 
covering about one-half of the lowermost row of dorsal scales 
on each side. The colors of the specimen in life were as fol- 
lows: Below, orange, fading anteriorly to a Capucine buff on 


labials and anterior chin shields, and becoming a little redder 


Occasional Papers of the Museum of Zoology 5 


at the posterior end of the body; under the tail, Brazil red; 
neck ring about scarlet, bordered posteriorly by a few specks 
of black; dorsal color behind neck ring, olive; head olivaceous 
black above. Total length, 291 millimeters; tail length, 59 mil- 


limeters. Sex, male. 


Diadophis amabilis vandenburgii, new subspecies 


Diagnosis: The distinctive features are: an uninterrupted, 
moderately wide neck ring (1% to 2% scales in width); 17 
rows of dorsal scales anteriorly (at least near the head), be- 
coming 15 posteriorly; the ventral color extending over from 
1% to 2 rows of dorsal scales; and the black spots on the 


belly unusually few and small. 


Type Specimen: California Academy of Sciences, number 


13748; collected by Joseph Slevin; June 20, 1907. 
Type Locality: Carmel, Monterey County, California. 
Range: Ventura County to Santa Cruz County, California. 


Description of Type Specimen: Ventral plates, 201; anal 
scute divided; caudal plates, 71; upper labials, 8; lower labials, 
8; preoculars, 2; postoculars, 2; temporals, 1+1-+-2 (labials, 
oculars, and temporals the same in number on each side) ; dor- 
sal scales in 17 rows, changing a little past the middle of the 
body to 15; neck ring about 1% scales wide, not interrupted ; 
ventral color covering about I 2/3 of the lowermost rows of 
dorsal scales. ‘The coloration in alcohol is about as follows: 
Above, brownish olive; head a little darker; neck ring, cinna- 
mon buff; beneath, pinkish-buff, a little lighter anteriorly, and 
a little more reddish posteriorly and under the tail; jower la- 
bials and chin spotted with black; on belly only a very few 


scattered black spots except for a line of them along each side 


6 University of Michigan 


on the ends of the ventrals, and one on the posterior end of 
most of the dorsal scales of the lowermost row on each side. 
Total length, 377 millimeters; tail length, 81 millimeters. Sex, 


male. 


Diadophis amabilis occidentalis, new subspecies 


Diagnosis: ‘The chief characteristics of this form are a rela- 
tively broad, uninterrupted neck ring, 1% to 2 scales in width, 
ventral color extending over 1% to 2 of the dorsal scale rows, 
flecks of black on the first two rows of dorsal scales, belly 
lightly spotted with black, and the dorsal scales in not 
more than 15 rows. ‘This form is intermediate between D. 
amabilis vandenburgti and D. amabilis pulchellus. From the 
former it differs in possessing not more than 15 rows of dorsal 
scales, in having a neck ring that averages slightly wider, and 
in having the ventral color extending on the average a little 
higher on the sides. From D. amabilis pulchellus it differs 
chiefly in having the light colored dorsal scale rows flecked 
with black; it also differs from this form in having on the 
average more black spots on the belly, a slightly narrower neck 


ring, and a lesser extent of ventral color on the sides. 


Type Specimen: Museum of Vertebrate Zoology, Univer- 
sity of California, number 7260; collected by H. E. Wilder, 
May 30, 1919. 

Type Locality: Bridgeville, Humboldt County, California. 


Range: From Sonoma County northward through Mendo- 


cine County, California, and perhaps to the Columbia River. 


Description of Type Specimen: Ventral plates, 197; anal 
plate divided; caudal scutes, 55; upper labials, 7; lower labials, ° 


8 on the left side and 7 on the right; preoculars, 2; postocu- 


Occasional Papers of the Museum of Zoology 5 


lars, 2; temporals, 1+1-++2 (upper labials, oculars, and tem- 
porals the same in number on each side); dorsal scales in 15 
rows throughout the body length; neck ring, 1% scales wide, 
not interrupted, but a slight median projection of its posterior 
black border present; ventral color covering about 1% of the 
lowermost rows of dorsal scales on each side. The coloration, 
as preserved in alcohol, is as follows: Above, about a dark 
olive-gray, darker posteriorly; head a little darker above; neck 
ring about cream color; ventral color a massicot yellow, be- 
coming reddish posteriorly and under the tail; ventral scales 
irregularly and rather prominently spotted with black, the 
light colored dorsal scales flecked with black, and the tail with 
only a very few small black spots. Total length, 442 milli- 


meters; tail length, 76 millimeters. Sex, female. 


Key TO THE WESTERN ForMs OF DIADOPHIS 


The ringneck snakes of the regalis and amabilis groups may 
nearly always be distinguished from all other forms of the 
genus in the United States by the possession usually of more 
than 180 ventral plates, and by the encroachment of the ventral 


color upon one or more of the lowermost rows of dorsal scales. 


Within these groups most specimens may be identified by 
the following key: 
a, Ventrals in males more than 206, in females more than 220. 


b, Neck ring present, 2 to 4 SEAMEG> iti eiyate Gel crest et ote a 8 oe wot oc 
D. regalis arizonae, subsp. nov. 


ote 6a wie a6 )e ss Nine @ Be Stare 6.0) @ are & 6 


(Central Arizona, south into Sonora.) 


b, Neck ring absent, or much reduced... 2... 2-1...) cence eevee eee 
Bae oe ee ie ces D. regalis regalis (Baird & Girard). 


(Central Texas to southeastern Arizona.) 
a, Ventrals in males less than 206, in females less than 220. 


c, Scale rows, 17-15 (rarely 15-15). 


University of Michigan 


d, Ventral color not covering more than 34 of the lowermost 
row of dorsal scales; belly usually conspicuously spotted 
with black. .D. amabilis modestus (Dumeril and Bocourt). 

(San Bernardino Mountains, Los Angeles County, and San- 
ta Catalina Island, California.) 


d, Ventral color covering from 1% to 2 of the lowermost rows 
of dorsal scales; belly usually only lightly spotted with 
Diack rere sc: eye D. amabilis vandenburgii, subsp. nov. 

(Ventura to Santa Cruz counties, California.) 


ce, Scale rows, 15-15 or 15-13 (rarely 17-15 or 15-17-15). 


e, Ventral color covering from % to % of the lowermost row 
of dorsal scales; neck ring only rarely interrupted; color 
above usually olive or bluish slate 


Meri aarehs 1S es ys D. amabilis similis, subsp. nov. 
(Southwestern San Bernardino County, California, south in- 
to the San Pedro Martir Mountains.) 


e, Ventral color covering usually more than % of the first 
row of dorsal scales. 


E Neck ring from 1 to 1% scales in width, often interrupt- 
ed; ventral color covering from % to 1% rows of dor- 
sal scales; belly well sprinkled with small black spots; 
dofsal color asually dark. 5 .-.02..2245.44... =o 
ee eee oe D. amabilis amabilis (Baird & Girard). 

(San Francisco Bay and the San Joaquin and Sacra- 
mento River valleys, California.) 


f, Neck ring from 1% to 3 scales wide, not interrupted; 
ventral color covering from 11%4 to 2 or more rows of 
dorsal scales; belly never heavily spotted with black. 


g, Two lowermost rows of dorsal scales flecked with 
black; belly rather conspicuously, although sparsely, 
marked with small black dots 
0 AN |. ae: D. amabilis occidentalis, subsp. nov. 

(Sonoma County north through Humboldt County, 
California, to the Columbia River.) 


g, Two lowermost rows of dorsal scales unicolor (not 
flecked with black); belly almost or quite unspotted. 
Sey es D, amabilis pulchellus (Baird and Girard). 

(Western slopes of the Sierra Nevada, south, perhaps, 
to Tejon Pass in California, and north to southern 
Oregon.) 


Occasional Papers of the Museum of Zoology 9 


ACKNOWLEDGMENTS 


This study would have been quite impossible without the 
generous cooperation of numerous individuals and institutions 
in the loaning of specimens, and the writer herewith takes 
pleasure in recording his indebtedness to Dr. Thomas Barbour 
of the Museum of Comparative Zoology, Mr. D. C. Davies of 
the Field Museum of Natural History, Mr. L. M. Klauber of 
the San Diego Zoological Garden, Dr. G. K. Noble of the 
American Museum of Natural History, Professor A. G. Ruth- 
ven of the Museum of Zoology of the University of Michigan, 
Professor J. O. Snyder of Stanford University, Dr. Leonhard 
Stejneger of the United States National Museum, Mr. Tracy 
I. Storer of the Museum of Vertebrate Zoology, Dr. John 
Van Denburgh of the California Academy of Sciences (for 
whom one of the new forms is named), Professor A. H. 
Wright of Cornell University, and Mr. L. EK. Wyman of the 


Los Angeles Museum of History, Science, and Art. 


al 


NUMBER 143 JULY , 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARzBor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


THE REPTILES OF THE DUTCH LEEWARD 
ISLANDS 


By ALEXANDER G. RUTHVEN 


The Museum of Zoology has recently received three collec- 
tions of reptiles from the Dutch Leeward Islands. In 1920, 
the University of Michigan-Williamson Expedition to Vene- 
zuela touched at Curacao and collected for a short time on 
that island. In 1922, Mr. M. A. Carriker, Jr., collected on 
Curacao for a few days and sent the reptiles to this Museum. 
The largest: collection was made by Dr. H. Burrington Baker,,. 
who was sent to Curacao, Klein-Curacao, Aruba, Bonaire, and 
Klein-Bonaire in the summer of 1922 to collect mollusks, rep- 
tiles and amphibians. 

These collections do not represent the entire fauna of the 
five islands, but they contain apparently new records, and 
several of the species are represented by large series which 
permit a more accurate description of the forms than 
has hitherto been possible. As the published lists (see 


bibliography) are evidently incomplete, and inaccurate owing 


2 University of Michigan 


to mistakes in the identification of the forms, the present list 
has been made to include the published records, even though 
there are many duplications. ‘The writer has had for study, 
through the courtesy of the Field Museum of Natural His- 
tory, some of the specimens referred to by Meek; but he has 


not been able to examine Cope’s specimens. 


List oF SPECIES 


Gonatodes albogularis (Duméril and Bibron). — Although 
Cope records this species both from Aruba and Curacao, and 
there are, according to Stejneger (1917), numerous specimens 
from Curacao in the United States National Museum, Baker 
secured but one specimen on Curacao. This is an old male and 
is quite typical of G. albogularis as distinct from G. fuscus 
and G. vittatus. 

Gonatodes vittatus (Lichtenstein). —Aruba (three specimens 
taken at Oranjestad). 

Cope lists the species from Aruba; and Boulenger (1885 a, 
p. 60) records a Curacao specimen in the British Museum un- 
der this name. Although carefully searched for elsewhere 
this species was only found in and about the town, where it 
was taken under rocks and in a house. 


Gymnodactylus antillensis (Van Lidth de Jeude).—Curacao 
(Willemstad; El Hato; Valley at Seroe Domi; Schaarlo; 
Campo Knip; Ronde Klip; Overzijde; Landhuis Knip; Tafel- 
berg of St. Hyronimus, 229.9 m.): Bonaire (hills of Santa 
Barbara, back of El] Hato de Bonaire; between Tanki Maraka 
and Porta Spafo; near Pos Frances; Seroe Wassau): Klein- 
Bonaire: Klein-Curacao. Some of the specimens of Van 
Lidth de Jeude were said to have come from Aruba, the others 
from Curacao. 


Occasional Papers of the Museum of Zoology 3° 


Common on all four andes rocks and in houses; a 


few were under the loose bark of trees. ) 


Phyllodactylus pulcher (Gray).—Curacao (Valley at Seroe 
Domi; Landhuis Knip; Schaarlo): Aruba (Sero Canashito ; 
Baranca Alto; near Malmok; near Oranjestad; near Perkien- 
tenboseh) : Bonaire (between Tanka Maraka and Porta Spajfio ; 
Santa Barbara beyond FE] Hato; Seroe Wassau): Klein-Bozi- 
aire. Recorded from Aruba by Cope, under the name P. ju- 


lient, and from Curacao by Werner and Van Lidth de Jeude. 


Not infrequent on Aruba, Bonaire and Klein-Bonaire, in the 
better wooded places; rare on Curacao; found under rocks 
and loose bark of dead trees. 

Boulenger (1887) has pointed out that P. martini Van 
Lidth de Jeude is synomymous with P. julieni Cope. Werner 
has listed a Phyllodactylus from Curacao as P. pulcher (type- 
locality tropical America), and Miss Joan Proctor has pro- 
nounced one of our specimens from Aruba identical with the 
type of P. pulcher. It is evident that the three names apply 
to one species which occurs on Aruba, Curacao, Bonaire, and 
Klein-Bonaire, and which should be known as Phyllodactylus 


pulcher (Gray). 


(Thecadactylus rapicaudus (Houttuyn).— Curacao (Campo 
Knip; Landhuis Knip): druba (Campo West Punt; Baranca 
Alto; Nooi Hundoe; near Perkietenboseh ) : Bonaire (between 
Tanki Maraka and Porta Spafio; Santa Barbara beyond El 
Hato di Bonaire). Listed from Aruba and Curacao by Van 
Lidth de Jeude, from Aruba by Cope, and from Curacao by 
Meek. 

Under the bark of trees and in buildings; sometimes under 


rocks at the base of trees. 


4 University of Michigan 
Anolis bonairensis, new species 


Diagnosis: ‘Tail moderately compressed, with serrated up- 
per edge. Dorsal scales small, those of the vertebral region 
larger than the laterals, smaller than the ventrals. Occipital 
scale large, larger than the ear opening, in contact with the 
scales of the supraorbital semicircles or separated from them 
by a row of granules, surrounded laterally and posteriorly by 
granules scarcely larger than the largest dorsals — at least 
much smaller than the scales of the forehead; scales of the 
supraorbital semicircles enlarged, separated or last one or two 
pairs in contact; supraocular scales enlarged, without keels. 
Tibia much shorter than the distance between end of snout 
and ear opening. 


Habitat: Bonaire and Klein-Bonaire, Dutch Leeward Is- 
lands. 


Type Specimen: Cat. No. 57221, Museum of Zoology, Uni- 
versity of Michigan; Seroe Grandi, 4% km. northeast of Kral- 
endijk, Bonaire; August 23, 1922; H. Burrington Baker, col- 
lector. 


Description of Type Specimen: Head moderate, about once 
and a half as long as broad, the length to the posterior border 
of the orbit equal to the length of the tibia; the hind limb ex- 
tended forward reaches the temporal region. Snout rather 
strongly depressed, forehead concave, no frontal ridges. Up- 
per head scales smooth, scales of the supraorbital semicircles 
large, last two pairs in contact with each other and the occipi- 
tal scale; about ten enlarged smooth supraoculars; loreal rows 
three ; occipital scale large, surrounded laterally and posterior- 
ly by granules scarcely larger than the enlarged dorsals, much 


smaller than those of the forehead; dorsal scales small, keeled, 


Occasional Papers of the Museum of Zoology 5 


those of the vertebral region larger than those on the sides: 
ventral scales larger than the dorsals, smooth. ‘Tail compress- 
ed, with serrated upper edge (regenerated). About 26 lam- 
ellae under the second and third phalanges of the fourth toe. 
Gular pouch rather large. Brown above with black crossbars 
bordered with yellow, and somewhat interrupted on the flanks ; 
ventral surfaces white, the sides of the abdominal region with 
faint spots, the chin mottled with dark brown, the gular pouch 
uniformly white (in alcohol). 


Heneth of head and body..:....... 66.5 mm. 
Bore ore med 0b es eee 19 mm. 
Eensth at tind ter 2200.0 232.2 50 =mm. 


Notes on Paratypes: The supraorbital semicircles in nine- 
teen specimens are entirely separated in six, in contact by one 
pair in ten and by two pairs in three. The enlarged supra- 
oculars are smooth in all specimens, and the occipital is sepa- 


rated from the supraorbitals by small scales in two specimens. 


The localities are as follows: Bonaire (Santa Barbara ; 
Montagne; Punta Blanco; Seroe \Wassau; Seroe Grandi): 
Klein-Bonaire. Usually (found upon trees. | Also recorded 


from Bonaire by Meek under the name 4. alligator. { Com- 


mon everywhere on trees and cliffs. 


Remarks: As is well known, anoles from a number of is- 
lands in the Carribean Sea have been referred to Anolis alli- 
gator. Barbour (1914, p. 281) has asserted that the Martin- 
ique form is distinct from the one on Trinidad, an assertion 
that has been supported by a study of specimens from both 
localities. A well marked difference is to be found in the size 
of the scales about the occipital plate. In a series of specimens 


these scales are as large as the scales of the forehead in those 


6 University of Michigan 


from Trinidad and much smaller, scarcely larger than the ver- 
tebral scales, at any rate much smaller than those of the fore- 


head, in those from Martinique. 


The Bonaire specimens agree with those from Martinique 
and differ from Trinidad specimens in having small scales 
about the occipital scale, and they differ from the Martinique 
form in having smooth supraoculars. and from both in having 
the supraorbital semicircles more widely separated. As stated 
above, in a series of nineteen specimens the semicircles are 
entirely separated in six, in contact by one pair in ten and by 
two pairs in three. In Martinique specimens and in some of 
those from Trinidad the enlarged supraoculars have a short 
keel, sometimes reduced to a low tubercle; and again in the 
Martinique and Trinidad specimens examined the semicircles 
are broadly in contact by three or four scales, the .forward 
point of contact being about on a level with, or a little posterior 
to, a line connecting the anterior margin of the first enlarged 


supraoculars. 


It is very probable that there are also differences in the col- 
oration of the three forms, but this cannot be satisfactorily 


determined with preserved material. 


It may be remarked here that the name* of the Martinique 
form is apparently Anolis roquet (Bonnaterre), Enc. Meth. 
Erpet., 1789, p. 54, pl. 9, fig. 5. This name is based on Le 
Roquet Lacépéde and antedates Merrem’s Anolis cepedii. That 
the Trinidad form is to be known as Anolis aeneus Gray (Cat. 
Liz., 1845, p. 205), and that this name is not available for the 
Bonaire form, seems to be established by the fact that the 
41 am indebted to Dr. Leonhard Stejneger for assistance with the 


nomenclature of these forms, and to Miss Mina Winslow for the 
examination of the type of Anolis aeneus Gray. 


Occasional Papers of the Museum of Zoology 7 


type of dA. aeneus has smooth supraoculars and large scales 
about the occipital plate: both characters distinguish it from 
Anolis roquet, and the second separates it from Anolis 


bonairensis . 


Anolis lineatus(Daudin).—Curacao( Valley at Seroe Domi; 
Willemstad; Tafelberg of Santa Barbara; El Hato; Seroe 
Salinja Abau; Schaarlo; Seroe Djerimi; Seroe Mansinga; 
Valley between Seroe Palomba and Seroe Baha Hoendoe; Hills 
south of Sint Willebrordus ; St. Christoffelberg, 500-1200 ft.) : 
Aruba (shore cliffs near Oranjestad; Seroe Canashito; Baran- 
ca Alto and Isla; near Boedoei; Hudishibana; near Perkieten- 
boseh). Listed from Aruba and Curacao by Meek and Van 
Lidth de Jeude, from Aruba by Cope, and from Curacao by 
Boulenger (1885 b, p. 39). Common everywhere on trees 
and bushes. 

A study of large series has failed to reveal any constant 


differences between the specimens from Curacao and Aruba. 


Iguana iguana (Linnaeus). — Curacao (observed, but not 
collected, on cliffs along the ocean at Campo Knip; on cliffs 
and nearby trees on Sint Christoffelberg, 1200 ft.; on cliffs 
south of Sint Willebrordus; and on cliffs and adjacent trees 
of the Tafelberg of Santa Barbara): Aruba (on a brush fence 
in Rooi Lamoenchi; on trees and in a sink-hole at Campo West 
Punt) : Bonaire (observed, but not collected, on cliffs of Seroe 
Wassau, and at Fontein of Bonaire). Listed by Cope, Meek, 
and Van Lidth de Jeude, from Aruba, and by Meek from 
Curacao. 

Ameiva bifrontata Cope. — Aruba (Oranjestad; Rooi La- 
moenchi; Boedoei; Campo West Punt; between Sint Nicolaas 
and Culebra; near Perkietenboseh; Seroe Canashito). Re- 


corded from Aruba by Cope. 


8 University of Michigan 


Common in all parts of the island but not as abundant as 
Cnemidophorus arubensis. 

Ten specimens are typical in scalation and show no tendency 
toward the subspecies divisus. The ground color is more uni- 
form and paler than in mainland series, black spots being ab- 
sent or very few in number. 


Cnemidophorus murinus (Laurenti)—Curacao (Schaarlo; 
Sint Willebrordus; Ronde Klip): Bonaire (Seroe Grandi, 
near Playa Makosji; Seroe Grandi, 414 km. northeast of Kral- 
endijk; Santa Barbara; Pos Frances; Seroe Wassau): Klein- 
Bonaire: Klein-Curacao. Listed from Curacao by Van Lidth 
de Jeude, Meek, and Cope, and from Bonaire by Meek. Cope’s 
C. murinus from Aruba is probably C. arubensis. . 

Abundant, according to Baker’s notes. 


Cnemidophorus arubensis (Van Lidth de Jeude). — Aruba 
(Oranjestad; Rooi Lamoenchi; Boedoei; Campo West Punt; 
near Perkietenboseh). 

Described from Aruba by Van de Lidth de Jeude. 

Abundant in all localities. 

This is Meek’s Cnemidophorus nigricolor, from Aruba; and 
it is probably Cope’s C. murinus from the same island. Al- 
though resembling C. murinus in coloration, C. arubensis is ap- — 
parently most closely related to C. lemmiscatus. It has eight 
rows of ventrals, as pointed out by Van Lidth de Jeude, and 
the subcaudal scales at the base of the tail are keeled as in 
C 1. lemniscatus and C. 1. gaigei. It differs from C. lemnis- 
catus in having smaller brachial and collar scales, and in color- 
ation. The pale spots are much larger and fewer in number 
in C. arubensis and the pale lines disappear, or are only faint- 
ly indicated, in old specimens. C. nigricolor from Los Roques 


is easily distinguished from C. arubensis by the color, by the 


Occasional Papers of the Museum of Zoology 9 


smooth scales beneath the base of the tail, and by the smaller 
brachial scales. 


Gymnophthalmus quadrilineatus (Linnaeus).—Curacao (1 
km. from Landhuis towards St. Christoffelberg ; St. Christoffel- 
berg, 1200 ft.; Landhuis Knip). Also listed by Van Lidth de 
Jeude from Curacao. 

Taken in grassy areas in thickets of acacias, and in humus 
among rocks. 


Tretioscincus bifasciatus (Duméril).—Aruba (one specimen 
in ant nest under rocks on hill between Rooi Spoki and Rooi 


Hundoe). Also recorded from Aruba by Cope. 


Dromicus antillensis (Schlegel). Recorded from Curacao 
by Van Lidth de Jeude. 


Leimadophis triscalis (Linnaeus ).—Curacao (a small spec- 
imen taken at Landhuis Knip). Also recorded from Curacao 


by Boulenger (1894, p. 129). 


Leptodeira annulata (Linnaeus).—Aruba (Campo Bubali; 
Mon Plaisir). Recorded from Aruba by Van Lidth de Jeude. 

The dorsal scale rows are 19-17-15 in six specimens. 

Found only in the matted clumps of branches of the older 
divi-divi trees. 

Crotalus terrificus (Laurenti). According to Baker, rattle- 
snakes are still to be found on Aruba. Meek has recorded a 
specimen, and Van Lidth de Jeude lists several under the name 


Crotalus horridus unicolor. 


IO 


IQI4. 


University of Michigan 


BIvLioCRAPHY 


Barsour, Tuomas. A Contribution to the Zoogeography of the 
West Indies, with especial Reference to Amphibians and 
Reptiles. Mem. Mus. Comp. Zool., XLIV, pp. 209-346. 


1885 (a) BouLeNcrr, G. A. Catalogue of the Lizards in the British 


Museum, I. 


1885 (b) Boutencer, G. A. Catalogue of the Lizards in the British 


1887. 


1894. 


IQI0. 


1917. 


1913. 


Museum, II. 
BouLeNncerR, G. A. The Zoological Record, XXIV, Rept. p. 9. 


Bovutencer, G. A. Catalogue of the Snakes in the British Mu- 
seum, II. 


Corr, E. D. Twelfth Contribution to the Herpetology of Trop- 
ical America. Proc. Amer. Phil. Soc., XXII, pp. 167-1094. 


LiptH DE JEupE, T. W. van. On a Collection of Reptiles and 
Fishes from the West Indies. Notes from the Leyden Mu- 
seum, IX, pp 120-139. 


Merk, S. E. Notes on Batrachians and Reptiles from the Is- 
lands north of Venezuela. Field Museum of Nat. Hist.,. 
Zool. Ser. VII, pp. 415-418. 


STEJNEGER, LEONHARD. Cuban Amphibians and Reptiles col- 
lected for the United States National Museum from 18909 
to 1902. Proc. U. S. Nat. Museum, LIII, pp. 2509-201. 


WERNER, FRANz. Neue oder Seltene Reptilien und Frésche 


des Naturhistorischen Museums in Hamburg. Mitt. Nat- 
urhist. Museum, XXX. 


“ 


NUMBER 144 DECEMBER 21, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arzor, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NOTE ON THE SPECIES OF EVERMANNICHTHYS, 
A GENUS OF SPONGE-INHABITING GOBIES 


By Cart, L. Hupss 


In 1917 Radcliffe* described a new species of sponge-inhab- 
iting goby from off the coast of North Carolina and from the 
west side of Florida. He named the species spongicola and 
referred it to the genus Garmannia. 


In 1921 Hubbs? based a new genus, Radcliffella, on Garman- 
nia spongicola, ‘There is no reason to doubt the propriety of 
generically distinguishing the species spongicola from Gobius 
paradoxus Gunther, which is the type of Garmannia Jordan 
and Evermann. 


In the meantime, however, Metzelaar (1919), in a paper only 
recently received by us, had named a very similar goby from 


Curacao Evermannichthys spongicola, new genus and species.’ 


* Radcliffe, Proc. U. S. Nat. Mus., 52, 1917, p. 423, fig. 

* Hubbs, Occ. Pap. Mus. Zool., Univ. Mich., No. 99, 1921, p. 2. 

*Metzelaar, Report on the fishes, collected by Dr. J. Boeke in the 
Dutch West-Indies 1904-1905, with comparative notes on the marine 


fishes of tropical West Africa, The Hague, 1919, p. 139, figs. 30, 40; 
Bijd. Dierk., 22, 1919, p. 141. 


2 University of Michigan 


Evermannichthys spongicola and Garmannia spongicola can- 
not be separated generically. Evermannichthys therefore 
replaces Radcliffella as the generic name for these gobies. The 
two species, however, appear from the original descriptions to 
be different, the spongicola of Metzelaar differing from the 
spongicola of Radcliffe in having a more slender body, shorter 


maxillary, and more rays in the second dorsal fin. 


The two species may therefore stand as follows.— 
1. Evermannichthys spongicola Radcliffe. 


2. Evermannichthys metzelaari Hubbs (new specific name). 


NUMBER 145 DECEMBER 21, 1923 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN Arsor,. MICHIGAN PUBLISHED BY THE UNIVERSITY 


TWO NEW FRESHWATER SNAILS FROM 
MICHIGAN 


By Mina L. WENSLOW 


The forms described below were found in the course of 
field work for the Museum of Zoology of the University of 
Michigan. Both were submitted to Dr. Bryant Walker, to 
whom grateful acknowledgment is made for helpful criticism. 
Tor a discussion of the subspecies of Planorbis antrosus 
reference may be made to Dr. Walker's “Notes on Planorbis 
II: P. bicarinatus,” in The Nautilus, Vol. XXIII, pp. 1-10, 
1909. It is interesting to find still another distinct form of 
P. antrosus in the same general northern region from which 
the varieties percarinatus, royalensis and portagensis have al- 
ready been listed. 


Planorbis antrosus jordanensis, new subspecies 
Figs. 1-5 
Shell with four and a, half closely coiled whorls, upper sur- 
ace decidedly flattened, concave, apical whorls deeply im- 


2 University of Michigan 


mersed, funicular; umbilicus narrow, deeply funicular; su- 
perior and basal carinae each forming a distinct, prominent, 
rounded cord; whorls flat, almost straight-sided; lines of 
growth strong, revolving sculpture distinct; aperture large, 
somewhat descending, auriculate, higher than wide, angled at 
the carinae; lip thickened within, edge sharp, somewhat ex- 
panded at the outer edge. Altitude, 6.5 mm.; diameter, 11.5 
mm. ; height of body whorl in front of aperture, 4.8 mm. 

Type Locality: South Arm of Pine Lake, about two miles 
north of East Jordan, Charlevoix County, Michigan. 

Type Specimen: Museum of Zoology, University of Mich- 
igan, No. 27440. Cotypes in the collection of Bryant Walker. 

This form combines features of at least two other varieties 
of Planorbis antrosus, but is typical of none of them. It is 
perhaps nearer to portagensis in size and general appearance, 
but differs from that form in having pronounced cords accent- 
ing the carinae, in flatter whorls, and in the wider angle 
which the upper edge of the aperture makes with the super- 
ior carina. It resembles royalensis, but differs from that sub- 
species in its smaller size, in the relatively smaller size and 
auriculate shape of the aperture, the two corded carinae, flat- 
ter whorls, and finer, more regular lines of growth. 

The series shows some variation in the degree of descent 
of the aperture, resulting in some specimens in the aperture 
being applied to the lower half only of the body whorl. In 
twenty adult specimens the altitude varies from 7.5 to 5.8 mm., 
diameter from 13.1 to 11.5 mm., and altitude of the body whorl 
in front of the aperture from 5.0 to 4.3 mm. 


Ferrissia michiganensis, new species 
Figs. 6, 7, 8 
Shell depressed, oval, slightly wider anteriorly, anterior and 
posterior margins broadly rounded, the latter slightly oblique 


Occasional Papers of the Museum of Zoology 3 


on the right side; right lateral margin slightly curved, left 
lateral margin more convex; the dorsal outline is flattened 
above posteriorly and slightly curved anteriorly, the greatest 
height being about in the centre of the shell, from which point 
it slopes slightly towards the apex and more rapidly towards 
the anterior margin, left lateral slope somewhat convex; the 
right lateral slope a little concave; apex blunt, slightly de- 
pressed, excentric, turned toward the right side, situated at 
about the posterior fourth of the length and about halfway 
between the median line and the right margin, radially striate ; 
lines of growth rather strong and irregular, the anterior slope 
is obsoletely radially rippled; light horn color. 

Length 4.25 mm., width 2.75 mm., altitude 1 mm. 

Type Locality: Willow Brook, west of Harbert, Chickaming 
Township, Berrien County, Michigan. 

Tvpe Specimen: Museum of Zoology, University of Mich- 
igan, No. 13057. Cotypes in the collection of Bryant Walker. 

This well marked species is the largest yet described from 
the Northern States of the depressed group of Ferrissias. 
While nearly as large as many examples of F. rivularis and 
F, tarda, the depressed form and apex readily differentiate it 
from either. 

Numerous examples were collected from dead leaves and 
sticks in a small pond above a dam in the brook, in 1917, and 
again in 1918 and 1922. In the same pond numerous other 
species of freshwater snails were found, among them the small 
Planorbis buchanensis Lea, Planorbis rubellus Sterki, Lym- 
naea columella Say, Lymnaea humilis modicella Say, and 
others. 


Figs. 
Fig. 


Fig. 


Figs. 


University of Michigan 


PLATE 1 


I, 2, 3. Planorbis antrosus jordanensis. Type. x 3. 


4. 


on 


Planorbis antrosus jordanensis. ‘Cotype, Mus. Zool., 
Univ. of Mich., No. 27442. Most inflated specimen. x 3. 


Planorbis antrosus jordanensis. Cotype, Mus. Zool., 
Univ. of Mich., No. 27441. Example of abruptly descend- 
ing aperture. x 3. 


6, 7, 8.. Ferrissia michiganensis. ‘Type. x 10. 


Piatr I 


New FRESHWATER SNAILS 


NUMBER 146 MAarcH 22, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ArRBorR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


TWO NEW SPECIES OF CISCO FROM THE 
GREAT LAKES? 


By WALTER KokELz 


Leucichthys alpenae, new species 


Argyrosomus prognathus Evermann and Smith, Rept. U. S. Comm. Fish. 
1894 (1806), p. 314-317 (in part). 

Leucichthys johannae Jordan and Evermann, Bull. Bur. Fish., 29, 1909 
(1911), p. 24-25 (in part). 

This form, currently known as the “longjaw”’, is one of the 
largest species of Leucichthys found in the Great Lakes. Indi- 
viduals not infrequently attain a length of 38 cm. (15 inches) 
and a weight of two pounds. The longjaws are well-flavored 
and moderately fat and are in demand by fish-smokers. The 
species occurs most abundantly at depths of less than 60 fath- 
oms and is generally distributed throughout Lake Michigan and 
Lake Huron, including Georgian Bay, where suitable condi- 
tions obtain. 


1These descriptions are published with the permission of the U. S. 
Commissioner of Fisheries. 


2 University of Michigan 


The type is a female specimen, to be deposited in the United 
States National Museum, 269 mm. in length to the base of the 
caudal, collected in Lake Michigan on June 15, 1923, 22 miles 
NNE of Charlevoix, Michigan, off Ie aux Galets, in 25-47 
fathoms of water. Paratypes, deposited in the Museum of 
Zoology of the University of Michigan, were obtained in Lake 
Michigan off the Michigan shore on August I1, 1920, 
i4 mi. SE 34°58, “and .on August” 12; “1920; ~15 sme 
SE x S % S of Manistique; on June 209, 1920, 
5 mi oN x H, on June 15, 1923, 22° mi;) NINE ace 
on August II, 1923, 3 mi. NW ¥% W of Charlevoix; on August 
10, 1923, 8 mi. NNW of Big Rock Point, and on August 21, 
1923, from an unknown location off Charlevoix; on June 22, 
1920, 5 mi. NNW, and on July 31, 1923, 5 mi. NW of Cat 
Head Light; on July 30, 1923, off the South Manitou Island; 
on October 4, 1920, 9 mi. north of Point Betsie; and on March 
20, 1919, I2 mi. west of Grand Haven. Other paratypes were 
taken off the Indiana shore on September 3, 1920, 22 mi. NW x 
N %N, on October 11, 1920, 20 mi. N x W 34 W, on Novem- 
ber 8, 1920, 18 mi. NNW and on November 19, 1920, 17 mi. 
NNW and 17% mi. NW x N 34 N of Michigan City ; and off 
the Wisconsin shore on September 23, 1920, 27 mi. ESE, of 
Milwaukee; on September 25, 1920, 18 mi. and also 5 mi. E % 
S of Port Washington ; on October 1, 1920, 11 mi. SE of Shey- 
boygan; on August 24, 1920, 10 mi. KE x N of Algoma; on 
August 23, 1920, 12 mi. EF, x S of the Sturgeon Bay Ship Chan- 
nel mouth; on August 18, 1920, 4 mi. west of Boyer ‘Bluff; 
and on August 19, 1920, 20 mi. EK % N of Rock Island. Speci- 
mens have also been taken in Lake Huron in Michigan waters 
off Cheyboygan, Rogers City, Alpena and Harbor Beach and in 
Canadian waters in Georgian Bay, off Lion’s Head and Wiar- 
ton; these are not designated as paratypes, and are not involved 
in the following description. 


Occasional Papers cf the Museum of Zoology 3 


The body is fusiform, somewhat compressed and elongate. 
The greatest depth through a point just in front of the dorsal 
comprises in the type 23% of the total length; in other adult 
specimens about 23-27%. The width is about 55% of the 
depth; in other specimens 50-55%. ‘The anterior dorsal profile 
of the body usually rises gradually from the occiput to the 
insertion of the dorsal, but it is sometimes somewhat steeper 
over its anterior half, particularly in the largest specimens. Be- 
hind the dorsal the line continues in a very faint curve to the 
caudal peduncle. ‘The ventral profile is rather strongly and 
uniformly curved from the tip of the snout to the caudal pe- 
duncle. The head, which is relatively short and deep, is con- 
tained 4.4 [(3.8) 4.1—4.4 (4.6) ]? times in the total length. 
In side view it is broadly triangular. Its dorsal profile is us- 
ually more or less faintly convex and forms a smooth arc 
continuous with that of the first half of the predorsal body 
contour. The premaxillaries are usually more or less pig- 
mented and are directed forward, ordinarily making an angle 
of 45°-60° with the horizontal axis of the head. The snout, 
seen from the side, is broad and rounded. The maxillary is non- 
pigmented and extends beyond the anterior edge of the pupil but 
seldom to its center. The eye is moderate in size, contained 4.6 
[ (3.8) 4.2—4.5 (5.2)] times in the head-length. The mandible 
is well developed and is usually longer than the upper jaw, 
though sometimes shorter. ‘The scales in the lateral line num- 
ber 75 [(71) 74—85 (96)]. The gill-rakers on the first 
branchial arch number 14+-25 [(11) 13 — 15 (17) + (20) 22 
—28 (30) = (33) 36—43 (46)]. The dorsal edge of the pec- 


toral is usually nearly straight. ‘The paired fins are relatively 


2These and succeeding figures in brackets are based on an examination 
of 289 paratypes ranging in length from 200 to 386 mm.; the usual as 
well as the extreme range in variation is given, the latter in parentheses. 


4 University of Michigan 


short. The pectorals are contained 2.2 [(1.6) 1.9—2.2 (2.5) ] 
times in the distance from their insertion to that of the ventrals ; 
the ventrals are contained 1.8 [(1.2) 1.4—1.7 (1.9)] times in 
the distance from their origin to that of the anal. 

In life the general appearance of the fish is silvery with a 
faint pink to purplish iridescence suffusing the sides. This 
reflection is strongest above the lateral line, becoming faint on 
the back and paling gradually toward the colorless belly. The 
color lying below the superficial iridescence is on the back a 
more or less intense pea-green to blue-green which loses its 
brightness toward the lateral line and becomes bright blue- 
green below it. On the back the color is obscured by a slaty 
cast due to fat in the epidermis and to the moderate, rather 
uniform pigmentation over the scales on the entire dorsal sur- 
face. The pigmentation is continued over the top of the head 
and extends also onto the preorbital area. It is likewise present 
but in diminished abundance on the sides of the body, chiefly 
above the lateral line. Four patches of green lie below the sur- 
face on each side of the mid-dorsal line of the head—three, the 
largest of which is rounded triangular, nearly contiguous, are 
located posterior to the center of the eye and extend backward 
to the occiput and the other, which is clubshaped, is located on 
the side of the mid-line, with its narrow end extending back- 
ward and inward to meet its companion of the other half of 
the head. A small patch of green also lies in the cartilage in 
front of the eye. The maxillaries, premaxillaries and mandible 
are whitish, all but the first usually showing at least some pig- 
ment. The cheeks and iris ‘are silvery with a trace of iridescence 
on the former and of bronze on the latter. The proximal one- 
third of the fins is often pale pinkish. The distal two-thirds is 
whitish except on the dorsal and caudal, on which it is more or 


less suffused with blackish. ‘The anterior border of the dorsal 


Occasional Papers of the Museum of Zoology 5 


and the lateral borders of the caudal are usually lined with 
black. The dorsal margin of the pectoral is often faintly 
sprinkled with black. Pigment dots are often present on the 
membranes connecting the longest rays of the anal and the 
ventrals occasionally also show pigment, but are usually im- 
maculate. 

Leucichthys reighardi,* new species 


L. reighardi is one of the smallest species of Leucichthys oc- 
curring in Lake Michigan, ranking in respect to size with kryi 
and hoyi. No specimens have been collected over 27 cm. in 
length and most of the specimens seen have been too small to 
gill in 234 inch nets. Like alpenae this species frequents depths 
of less than 60 fathoms and is taken with alpenae and the other 
deep-water species of Leucichthys for the smoked fish trade. 

The type is a female specimen, to be deposited in the 
United States National Museum, 210 mm. in length to the 
base of the caudal, collected in Lake Michigan on 
April 1, 1921, 18 miles N x W of Michigan City, 
Indiana, at a depth of 30-35 fathoms. Paratypes, de- 
posited in the Museum of Zoology, were obtained in 
Lake Michigan off the Michigan shore on March 20, 1919, 12 
mi. west of Grand Haven; off the Indiana shore on September 
3, 1920, 22 mi. NW x N % N, on October 11, 1920, 20 mi. 
N x W 3% W, on November 8, 1920, 18 mi. NNW, on No- 
vember 19, 1920, 17 mi. NNW, on March 2, 1921, 14 mi. 
NNW, on March 4, 1921, 15 mi. NNW, and on April 1, 1921, 
18 mi. N x W of Michigan City. Other paratypes were 
’ taken off the Wisconsin shore on March 24, 1919, 

m an unknown location off Milwaukee, and on Sep- 
tember 23, 1920, 27 mi. ESE, of Milwaukee; on September 25, 


3Named in honor of Professor Jacob Reighard, Department of 
Zoology, University of Michigan. 


6 University of Michigan 


1920, 18 mi. EF % S and on May 26, 1922, 8 mi. NE of Port 
Washington; on September 28, 1920, 5 mi. and 40 mi. SE x E 
and on October 1, 1920, 11 mi. SE of Sheyboygan; on August 
24, 1920, 10 mi. E x N of Algoma; on August 23, 1920, 12 mi. 
Ex S of the Sturgeon Bay Ship Channel mouth; on August 18, 
1920, 3 to 5 mi. WNW of Boyer Bluff, and on August 
19, 1920, 20 mi. F % N of Rock Island. Other specimens, not 
designated as paratypes, and not involved in the description of 
the species, were collected in Lake Michigan off the Michigan 
shore on August 12, 1920, 15 mi. SE x S % S of Manistique; 
on June 29, 1920, 5 mi. N x E, on June 30, 1920, 3 mi. NW, on 
June 15, 1923, 22 mi. NNE.and on August 11, 1923, 3 mi. NW 
% W of Charlevoix ; on August 10, 1923, 8 mi. NNW of Big 
Rock Point, and on August 21, 1923, from an unknown loca- 
tion off Charlevoix; on June 22, 1920, 5 mi. NNW and on 
July 31, 1923, 5 mi. NW of Cat Head Point; on June 23, 1920, 
off Northport Point; on October 4, 1920, 9 mi. north of Point 
Betsie ; on August 27, 1920, 4 mi. west, and on August 28, 1920, 
g mi. NW of Manistee. Still others were collected in Lake 
Ontario, in New York waters off Wilson, Sodus Point, Os- 
wego, Selkirk and Sandy Pond, and in Canadian waters off 
Brighton; in Lake Superior off Grand Marais, Minnesota, and 
at several locations on the Ontario shore between Fort William 
and Rossport, and also in Lake Nipigon. 

The body is little compressed, much less than in any other 
member of the genus, excepting artedi, and as seen from the 
side, tapers smoothly and regularly to the head and tail from 
the deepest portion of the body which is through a point at the 
front of the dorsal. This dimension is in the type 23%, in most 
of the other adult specimens at hand 22-26%, of the length. 
Occasionally an individual is taken in which this figure rises to 


29% and in such specimens the predorsal body profile is steeper 


Occasional Papers of the Museum of Zoology 7 


over its anterior half. The width of the type specimen is 62% 
of the depth; in other specimens 55-65%. ‘The head is short 
and is contained 4.4 [(3.9) 4.2—4.4 (4.8) ]* times in the total 
length. It is of moderate depth, bluntly triangular in side view; 
its dorsal contour, not including the premaxillaries, straight or 
faintly convex. ‘The snout is likewise short, truncated in side 
view on account of the nearly vertical direction of the premaxil- 
laries, and is contained 3.8 [(3.5) 3.6—4.0 (4.4) ] times in the 
head. The premaxillaries are always heavily pigmented, and 
usually make an angle of 60°—70° with the horizontal axis of 
the head, so that their tip is usually at or below the lower edge 
of the pupil. The maxillary is always more or less pigmented, 
the cutting edge usually rimmed with black halfway to its distal 
end; it is short, being contained 2.7 [(2.5) 2.6—2.8 (3.0)] 
times in the head. The eye is large, is contained 3.8 [(3.6) 3.9 
—4.2 (4.4)] times in the head and is situated in the second 
quarter of the head-length, encroaching more or less on the 
third. The lower jaw is always shorter than the upper and is 
usually heavily tipped with black. The lateral line scales num- 
ber 72 [(67) 72—81 (96)]. ‘The gill-rakers on the first 
branchial arch number 14-++-23 [(11) 12—14 (16) + (20) 21 
—24 (27) = (31) 34-38 (43)]. The paired fins are rela- 
tively short. The pectoral length divided into the interval be- 
tween the pectoral and ventral insertions equals 2.1 [ (1.8) 2.0— 
2.5 (2.8)]; the ventral length into the distance from its origin 
to the anal equals 1.4 [(1.2) 1.4—1.7 (1.8) ]. 

The color of living paratypes has not been recorded but it is 
probably not different from that of the race of reighardi oc- 
curring in the northern basin of Lake Michigan. As in other 


4These and succeeding figures given in brackets are based on an ex- 
amination of 145 paratypes ranging in length from 200-257 mm.; the 
usual as well as the extreme range in variation is given, the latter in 
parentheses. 


8 University of Michigan 


species of Leucichthys the general tone is silvery and individuals 
are taken whose color differs in no material way, excepting 
heavier pigmentation, from specimens of alpenae just described. 
Many examples, however, show very little greenish color on 
the back. In such individuals the cranial patches are faint pea- 
green and the cranial cartilages and the fin bases have a sepia 
tone. The back is of a very pale blue-grey tint with traces of 
pea-green beneath. The iridescence of the sides is pinkish. 


NUMBER 147 Aprin 26, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NEW LEPOSOMA FROM PANAMA 
By A. G. RuTHVEN AND HELEN T. GAIGE 


A number of specimens of a Leposoma which seems to differ 
widely from the hitherto known species, were collected by the 
Bryant Walker Expedition to Panama (1923). We take great 
pleasure in dedicating this form to Dr. John Grant South, 
United States Minister to the Republic of Panama. 


Leposoma southi, new species 


Diagnosis: Head scales very rough, longitudinally striated; 
two frontonasals; praefrontals half the length of the frontal 
and smaller than the frontoparietals; three pairs of chin 
shields, the two anterior pairs in contact; scales from the eye 
to shoulder very rough (smoother in dispar and taeniata) .* 

Type Specimen: Cat. No. 48065, Museum of Zoology, Uni- 
versity of Michigan. Progreso (90 ft. elev.), Chiriqui Prov- 
ince, Panama; April 19, 1923; H. T. Gaige, collector. 

Description of Type Specimen: Form of head and body as 
in L. dispar; head scales very rugose, strongly striated; two 
frontonasals, each slightly concave behind; a pair of prae- 


1 We are indebted to Dr. G. K. Noble, American Museum, for the 
opportunity of examining a paratype of L. taeniata. 


2 University of Michigan 


frontals, each half the length of the frontal, smaller than the 
frontoparietals; parietals almost as wide as the interparietal ; 
four supraorbitals; two frenoorbitals; six upper, five lower 
labials; one anterior chin shield, followed by three pairs, the 
sides of the anterior two pairs in contact, the posterior pair 
much smaller, separated by a group of three small scales, and 
bordered behind by a row of enlarged scales which are sepa- 
rated by several smaller scales; a series of granules across the 
throat from ear to ear; scales from eye to shoulder very rough, 
almost tubercular; scales on the throat narrower and more 
pointed than the ventrals; ventrals strongly keeled, mucro- 
nate; anal plates five; scales around the middle of body 23; 
scales from occiput to base of tail 31; scales from chin granules 
to vent 33. 

Snout to vent 32 mm.; snout to ear 7 mm.; snout to eye 
3 mm.; greatest width of head 5 mm. 

Color in life: head and tail deep brown, almost black; a 
stripe of reddish brown from occiput to tail, becoming lighter 
on the sides, where it is bordered by a dark brown stripe; 
below reddish yellow; three light spots on upper lip and five 
black spots on lower lip. 

Remarks: There are nine paratypes of the new species in 
the collection which vary little in color and scalation. It 
should be noted that the smallest specimen has a deep groove 
down the center of the frontonasal, but it is not actually 
divided. The scale counts of eight specimens average: around 
the middle of the body 2414, from occiput to base of tail 32, 
and from the chin granules to vent 34. The main variation 
is in the arrangement of the chin shields. The third pair 
may be separated by one, or as in the type, by three small 
scales bordered behind by scales of equal size, or by a larger 
scale which may be bordered behind by two scales of the same 
size with still smaller ones behind, or the smaller, irregular 
scales may be in direct contact with the larger scale. 

The Museum of Comparative Zoology has sent us for com- 
parison a Leposoma (collected at Suretka, Sixaola River, 
Costa Rica, by E. R. Dunn) that corresponds fairly well with 


Occasional Papers of the Museum of Zoology 3 


LL. southi with the exception of the chin shields. The shields 
of the third pair are small, separated by two scales almost as 
large, and behind them is a row of subequal enlarged scales. 

Our specimens were all collected in the ground litter on the 
forest floor in the District of Alanje. 


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NuMBER 148 ApRIL 26, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


STUDIES OF THE FISHES OF THE ORDER 
CYPRINODONTES 


V. Nores on SPECIES OF GOODEA AND SKIFFIA 


By Cari L. Husss 


i 


The species of Goodea and Skiffia belong to a peculiar group 
of viviparous eyprinodonts, the family Goodeidae, which has 
been discussed in the first paper of this series. The fishes of 
this family form a large element in the highly distinctive 
Lerma River fauna of Mexico.* 


II 


GENUS GOODEA JORDAN 


Goodea Jordan, Proc. U. S. Nat. Mus., 2, 1879, p. 299; Jordan and 
Evermann, Bull. U. S. Nat. Mus., 47, pt. 1, 1896, p. 685; Meek, Publ. 
Field Mus., Zool., 3, 1902, p. 100; ibid., 5, 1904, p. 136; Regan, Biol. 
Cent.-Am., Pisces, 1908, p. 90; Ann. Mag. Nat. Hist., (8)7, 1911, 
p. 325.—Haplotype, Goodea atripinnis Jordan. 

Xenendum Jordan and Snyder, Bull. U. S. Fish Comm., 19, 1899 
(1900), p. 127.—Orthotype, Xenendum caliente Jordan and Snyder = 
Goodea atripinnis Jordan. 


1See Meek, Publ. Field Mus., Zool., 5, 1904, pp. xxxvii-lii, 109-124, 
136-144. 


LO 


University of Michigan 


The identity of Xenendum with Goodea has already been 
indicated by Meek. 

It may be of some historic interest to mention here that 
the viviparity of Goodea was noted as early as 1896, by Dr. 
Herrera.” 


1. Goodea luitpoldi Steindachner 


Characodon luitpoldi Steindachner, Denk. Akad. Wiss. Wien, 42, 1895, 
p. 528, pl. 2, fig. 3; Jordan and Evermann, Bull. U. S. Nat. Mus., 47, 
pt. 3, 1898, p. 2832; Pellegrin, Bull. Mus. Hist. Nat. Paris, 7, 1901, p. 205. 

Goodea luitpoldi Meek, Publ. Field Mus., Zool., 3, 1902, p. 101, pls. 
22-24; ibid., 5, 1904, p. 139, fig. 42, pl. 1-2. 

Characodon (Goodea) atripinnis Herrera, Cat. Col. Peces, Mus. Nae., 
Mexico, 1896, p. 31 (not of Jordan). 

Goodea atripinnis Regan, Biol. Centr.-Am., Pisces, 1908, p. 91; Ann. 
Mag. Nat. Hist., (8)7, 1911, p. 235, pl. 8 (not of Jordan). 

(2) Goodea atripinnis Fowler, Proc. Acad. Nat. Sci. Phila., 68, 1916, 
p. 432 (?not of Jordan). 

Xenendum «xaliscone Jordan and Snyder, Bull. U. 8. Fish Comm., 19, 
1899 (1900), p. 128, fig. 9; Jordan and Evermann, Bull. U. 8S. Nat. Mus., 
47, pt. 4, 1900, p. 3153. : 


Specimens from Lake Chapala (‘‘Xenendum saliscone’’) 
have a band of fine teeth behind the main row, as in topotypes 
of luitpoldi from Lago de Patzcuaro, and agree also in other 
respects. 

Goodea luwitpoldi is a species of the larger lakes. It is not 
known from the smaller streams, which are inhabited by 
Goodea atripinnis. Specimens of the latter form from Lake 
Cuitzeo, in fact, show an approach, in some respects, toward 
luitpoldi. 

The present species is a larger fish than atripinnis, and 
usually differs from it in the more anterior position of the 
dorsal fin, and constantly in the smaller size of the seales. 

The young of lwitpoldi are blotched like those of atripinnis. 
The blotches soon become obsolete, however, the color pattern 
of the adult of both sexes consisting of longitudinal streaks 
between the scale rows. 


2 Cat. Col. Peces, Mus. Nac., Mexico, 1896, p. 31. 


Occasional Papers of the Museum of Zoology 3 


2. Goodea atripinnis Jordan 


Goodea atripinnis Jordan, Proc. U. 8. Nat. Mus., 2, 1879, p. 299; 
Bean, ibid., 21, 1898, p. 541; Jordan and Evermann, Bull. U. S. Nat. Mus., 
47, pt. 1, 1896, p. 685; Meek, Publ. Field Mus., Zool., 3, 1902, p. 100; 
ibid., 5, 1904, p. 140, fig. 48; ibid., 7, 1907, p. 156 (in part: specimens 
from San Miguel only). 

Characodon variatus Woolman, Bull. U. S. Fish Comm., 14, 1894, p. 
62 (in part). 

Xenendum caliente Jordan and Snyder, Bull. U. S. Fish Comm., 19, 
1899 (1900), p. 127, fig. 8; Jordan and Evermann, Bull, U. S. Nat. 
Mus., 47, pt. 4, 1900, p. 3152. 

Goodea calientis Meek, Publ. Field Mus., Zool., 3, 1902, p. 100; Regan, 
Biol. Centr.-Am., Pisces, 1908, p. 91. 


The material described and recorded by Jordan, Woolman, 
Jordan and Snyder, and by Meek, has been re-examined. The 
specimens recorded by Woolman from Salamanca as Chara- 
codon variatus represent the females of both that species and 
of Goodea atripinnis. Having identified specimens of the 
present species with Characodon variatus, it is not surprising 
that he was unable to satisfy himself of the identity of Chara- 
codon variatus (male) with C. ferrugineus (female, of 
variatus ). 

Goodea atripinnis is essentially a stream species. It is not 
represented in the collections from lakes Chapala, Patzcuaro 
and Zirahuen which have been examined. All of the speci- 
mens from these three lakes represent the larger, finer scaled 
species, Goodea luitpoldi. It is highly probable that all 
records of this species from Lake Patzcuaro refer to lwitpoldi. 

The only lake specimens of atripinnis seen are from the 
small Lake Cuitzeo.2 In addition to their lighter color, they 
have the dorsal fin farther forward than usual. The distance 
from the origin of the dorsal to the end of the caudal when 
measured forward extends nearly to the eye, instead of little 
beyond the opercular margin, as usual in the several series of 

3In recording these specimens Meek (1900) mentioned that ‘‘the 
specimens from Lake Cuitzeo are very light in color, a feature character- 
istic of all the fishes taken from this and P&tzcuaro Lake.’’ Regan 


(1908) erroneously construed this sentence to mean that specimens of 
atripinnis were being recorded from Patzcuaro. 


4 University of Michigan 


atripinnis at hand (for example, the topotypes of X. caliente). 
In other lots, however, for instance that from San Miguel, the 
dorsal may occupy either position, or an intermediate one. 
Provisionally these differences are interpreted as of only 
racial, not subspecific, value. 

As noted by Dr. Meek (1902), the young are blotched, in 
coloration resembling Zoogoneticus robustus and the young of 
Characodon variatus, species of other genera (but of the same 
family). These brown blotches are usually retained more or 
less distinctly by the female, but soon become obsolete in the 
male, which is the more deeply colored. In the adult the 
color pattern, exclusive of the brown blotches on the female, 
consists of dark marks, one on each seale. Each of these is 
usually wedge-shaped, but not infrequently becomes narrowed 
dorsoventrally and bordered with a very fine lighter streak. 
In other specimens the dark pigment is mostly concentrated 
toward the base and upper and lower angles of the scales, 
leaving the center whitish, as in G. lwitpoldi. The vertical 
fins are often black in either sex. 


3. Goodea captiva Hubbs, new species 


Goodea atripinnis Meek, Publ. Field Mus., Zool., 7, 1907, p. 156 (in 
part: specimens from Jesus Maria only). 

This species is closely related to Goodea atripinnis, from 
which it has doubtless been derived by isolation following 
stream capture. The type-specimens come from one of those 
tributaries of the Rio Panuco which are known to have ex- 
tended their course backward until they have drained what 
was formerly a part of the Lerma System. Another set of 
specimens from the Panuco Basin, namely that from San Juan 
del Rio, remains typical of G. atripinnis. Goodea toweri, also - 
from the tributaries of the Rio Panuco, is a very different 
species. 

Goodea captiva differs from G. atripinnis in the form of the 
body, the contours being more arched, the caudal peduncle 
more slender; in the more anterior position of the dorsal fin, 
the origin of which is equidistant from end of caudal and a 


Occasional Papers of the Museum of Zoology 5 


point near the front margin of the eye, rather than some point 
in the postorbital part of the head; in coloration; on the 
average in the larger scales, these being in 32 to 36 instead of 
34 to 38 transverse series; and in the shorter gill-rakers, few 
more than half as numerous as in atripinnis. 

Holotype—Taken by Dr. 8. E. Meek, together with the 
numerous paratypes, in an upper tributary of the Rio Panuco, 
at Jesus Maria, Mexico; an adult male 46 mm. long to caudal 
base ; Cat. No. 5557, Field Museum. Regarding this collection, 
Dr. Meek (I. c., p. 153) remarks: ‘‘There is a small stream at 
Jesus Maria which belongs to the Rio Panuco system. It is 
almost without water during the dry season. At the Hacienda 
a dam is built across the narrow valley forming above it a 
small lake. In this 4 species of fishes were taken,’’ including 
the types of G. captiva. The following description is based 
upon the type, 46 mm. long to caudal. base, and is supple- 
mented by measurements and counts of five males 40 to 45 mm. 
long, and of five females, 45 to 49 mm. long. 

The body is deeper and with more arched contours than in 
G. atripinnis; the dorsal contour is slightly elevated at the 
occiput and gradually curved thence to the origin of the 
dorsal; the ventral contour is deeply curved—more deeply in 
the female than in the male—from the mouth to the anus, the 
deepest point in the curve being before the front of ventrals; 
greatest depth, 2.55 (2.5 to 2.7 in adult male paratypes; 2.6 to 
2.8 in adult female paratypes; in young specimens of equal 
size, the males are a little deeper than the females). From 
the origins of the dorsal and anal fins the contours of the male 
converge to the slender caudal peduncle so abruptly that if 
produced they would meet at an angle of nearly ninety degrees 
(the contours converge less rapidly in the female, as in that 
sex the greatest depth is farther forward than in the male; 
the distance between the origin of the dorsal and of the anal 
fins is about the diameter of the eye greater in the adult male 
than in the adult female). Least depth of the caudal 
peduncle, a little more than half its length behind anal fin; its 
length about equal to length of head. 


6 University of Michigan 


Head deep and heavy, the contour straight from occiput to 
tip of snout. Length of head, 3.35 (3.35 to 3.5, in male para- 
types; 3.65 to 3.85 in females) ; length of snout, 3.4 (3.4 to 
3.6, males; 3.3 to 3.5, females) ; length of eye, 3.6 (5.4 to 3.7, 
males; 3.6 to 4.0, females); width of slightly convex inter- 
orbital, 2.3 (2.2 to 2.4, each sex). Mouth wide, transverse, 
the width of the cleft about equal to length of snout; mandible 
heavy, but flexible; broadly projecting beyond the premaxil- 
laries. Teeth of two kinds, an outer row of incisors and an 
inner series of small villiform teeth. The incisors are dilated 
and bifid distally, and are movable; as in atripinnis, the teeth 
of the outer row are crowded into a biserial arrangement, 
alternating in their insertion. Gill-rakers short and flat, the 
longest only one-third as long as the eye; 24 to 26 gill-rakers 
on the first arch (40 to 45 in atripinnis). Intestines long and 
much convoluted. Viviparous, the largest females containing 
small embryos. 

Fins all of decidedly greater expanse in the male than in 
the female. The dorsal especially broad and elevated in the 
male, reaching when depressed to the first of the procurrent 
caudal rays; in the female reaching little more than two-thirds 
as far. Height of anal more than half length of head in male, 
less than half head in female; length of pectoral 1.4 in head 
in male, 1.6 in female; pelvic reaching anus in male; not so 
far in female. Fin-rays: dorsal, 13; anal, 14 or 15 (in each 
sex) ; the first 6 anal rays unbranched and shortened in the 
male. 

Seales large, in 35 (32 to 36, numerous paratypes) series 
from above branchial aperture to caudal base (34 to 38, in 
atripinnis ; 39 or more in luitpoldi) ; in 14 (13 to 16) rows 
from pelvic base up to mid-dorsal line. 

Color of type (adult male) : very dark; sides of body black- 
ish brown, the dark color extending farthest downward above 
anus; the caudal peduncle, black; scale margins everywhere 
light; back of trunk lighter; belly and edges of caudal 
peduncle, light ; top of head black ; upper half of opercle black, 
lower half silvery. Dorsal fin blackish at base, grading to 


Occasional Papers of the Museum of Zoology 7 


clear distally; caudal fin dusky at base, shading to black 
posteriorly, but with an abrupt, clear margin; lower fins clear. 

The development of the color pattern in the two sexes offers 
some points of interest. Young males 23 to 26 mm. long are 
colored very much like the young females. The body is 
marked with rather small black spots of various shapes, mostly 
absent along light streaks just above and just below an irregu- 
lar axial dark stripe. In males of 30 mm., the spots have 
become obsolete, or rendered indistinct by the deepening of 
the ground color. Even in the half grown, however, the 
widened and deepened axial band remains; there is also fre- 
quently evident a dark blotch between this streak and the 
anus. Females less than 30 mm. long have the rather faint 
axial streak intensified by the concentration along it, and the 
partial absence just above and below it, of the small black 
irregular spots which seatteringly cover the body. In the 
females over 30 mm. long the spots are smaller, and in those 
over 35 mm. long they have become indistinct. In females 
over 45 mm. long the lateral band is also very indistinet or 
not evident, the pattern consisting of the cross-hatching caused 
by the disposition of pigment about the margins of the scales. 
The coloration of the adult female has thus come to resemble 
that of the adult male, but by a slower rate of change. The 
color of the body is less intense in the female than in the 
male, however, in all specimens over 28 mm. long. The dorsal 
and caudal fins become dusky, but not blackish as in the male. 


Et 


GENUS SKIFFIA MEEK 


Skiffia Meek, Publ. Field Mus., Zool., 3, 1902, p. 102; ibid., 5, 1904, 
p. 141. 


The species referred to Skiffia differ from those of Goodea 
chiefly in the more advanced location of the dorsal fin. 


4. Skiffia bilineata Bean 


Characodon bilineatus Bean, Proce. U. S. Nat. Mus., 10, 1887, p. 371, 
pl. 20, fig. 2; Jordan and Evermann, Bull. U. S. Nat. Mus., 47, pt. 1, 
1896, p. 668; pt. 4, 1900, pl. 119, fig. 293. 


§ University of Michigan 


Skiffia bilineatus Meek, Publ. Field Mus., Zool., 3, 1902, p. 105. 
Skiffia bilineata Meek, Pub. Field Mus., Zool., 5, 1904, p. 144, fig. 45. 
Goodea bilineata Regan, Biol. Centr.-Am., Pisces, 1908, p. 92. 

Sexual dimorphism has been carried to an extreme in this 
well-marked species. Four males, 19 to 22 mm. long to the 
caudal fin, collected by Meek at Huingo, Lake Cuitzeo, Mexico, 
have been examined. In these the body is more slender than 
in the female, and the vertical fins are much larger. The black 
dorsal fin reaches to the base of the upper caudal rays. The 
highly characteristic color pattern of the female is not devel- 
oped in the male, only the anterior portion of the main median 
streak being apparent. In contrast the male has 12 to 20 
spindle-shaped vertical bars, of greatly varying height, dis- 
tributed along a line occupied by the lateral band in the 
female. In both sexes the edges of the caudal peduncle are 
black. 


5. Skiffia lermae Meek 


Skiffia lermae Meek, Pub. Field Mus., Zool., 3, 1902, p. 102, pl. 25; 
ibid., 5, 1904, p. 142. pl. 8. 

Goodea lermae Regan, Biol. Centr.-Am., Pisces, 1908, p. 92. 

(?)Skiffia variegata Meek, Publ. Field Mus., Zool., 3, 1902, p. 104, 
pl. 25; ibid., 5, 1904, p. 143, fig. 44. 

The types of Skiffia variegata, from Lake Zirahuen, are 
similar in form to specimens of like size from Lake Patzeuaro 
(lermae). They have the same number of fin rays (usually 
13 in both dorsal and anal fins) and the same number of seales 
(usually 35 or 36 rows). 

The differences which are apparent involve only the colora- 
tion. The spots are larger in those from Zirahuen (variegata) 
than in those from Patzeuaro (lermae), but the spot on the 
base of the caudal rays is not vertically elongate as in those 
from Patzcuaro. Those from Chaleo resemble ‘‘variegata’’ in 
coloration, while those from Celaya are like typical lermae. 

It remains to be determined, however, whether these color 
differences are of taxonomic significance. 


NuMBER 149 AprIL 26, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


DESCRIPTION OF AN AMEIVA FROM TESTIGOS 
ISLAND, VENEZUELA 


By ALEXANDER G. RUTHVEN 


A specimen of Ameiva, collected on Testigos Island, was 
recently sent to the writer for study from the Museum of 
Comparative Zoology. The specimen differs in several char- 
acters from its nearest relative—A. bifrontata Cope. The 
differences are not great, and only one specimen is at hand, 
but, since the characters are distinct and the genus has ap- 
parently not been recorded from the island, it seems proper 
to describe the form as new. 


Ameiva insulana, new species 


Diagnosis: Ten longitudinal rows of ventral plates ; arrange- 
ment of head scales as in Ameiva bifrontata; caudal scales 
straight keeled; enlarged gulars in a band across the throat, 
last three supraoculars together surrounded by granules; 
outer toe reaching nearly as far as inner; brachials and ante- 
brachials not continuous; postbrachials slightly enlarged; 
hind leg reaching to the anterior border of the ear. 

Type Specimen: No. 14025, Museum of Comparative Zool- 
ogy ; Testigos Island. 


2 Unversity of Michigan 


Description. of Type Specimen: Nostril between the nasals; 
five occipitals, the anterior margin of the middle one as wide 
as or wider than the adjacent ones; frontal divided trans- 
versely, with low longitudinal keels; second and third supra- 
oculars large, the first small, the fourth very small, the last 
three entirely surrounded by granules; six superciliaries; 
loreal undivided; five and six supralabials to below middle of 
eye; two pairs of chin shields in contact on the median line; 
mesoptychial scales about the size of the largest granules. 
Dorsal seales small; ventrals in ten longitudinal rows; trans- 
verse rows about thirty-four; preanal plates small, four sub- 
equal scutes in a row posteriorly. One row of large brachial 
plates not continuous with the antebrachials, which are in one 
row. <A group of slightly enlarged rhomboidal postbrachials. 
Femoral plates numerous; tibial plates in three rows. Femoral 
pores fifteen. Tip of longest toe reaching the anterior margin 
of the ear. 

The color is faded but evidently the ground color was oliva- 
ceous, and the pattern consisted of light stripes and dark spots 
arranged as follows: A pale median stripe obsolete on the 
neck; an upper narrow lateral stripe continuous from the 
head and extending on the tail; a lower narrow lateral stripe 
continuous from the ear to the shoulder, broken up into spots 
behind the shoulder for half the body length, and again con- 
tinuous from the middle of the body and on the tail; a series 
of irregular dark spots between the lateral stripes; two rows 
of regularly triangular and alternately placed spots on the 
back, separated by the median pale stripe; an irregular yellow 
stripe on the posterior side of the thighs. 


mm. mm 
otal length oy.ce csc 205 Length of hind leg ........... 47 
Length of head ................. 18 ‘Lieneth oftath c.cs)22 144 


Remarks: This species is evidently very close to Ameiva 
bifrontata Cope as it exists on Aruba. If the specimen is 
typical, the principal differences are in the wider median 


Occasional Papers of the Museum of Zoology 3 


occipital plate, the smaller postbrachials, the discontinuity be- 
tween the enlarged brachials and antebrachials, the smaller 
preanals, and the longer hind leg in the Testigos form. 
Slight as are the differences, it seems best to give the Testigos 
Ameiva specific rather than subspecific rank because of its 
isolation. 


NuMBER 150 AprIL 26, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


A NEW SNAKE OF THE GENUS ARIZONA? 
By FRANK N. BLANCHARD 


It has been apparent to the writer for some time that the 
snakes long referred to Arizona elegans comprise two races 
sufficiently well-marked in appearance and distribution to be 
designated by separate names. Kennicott’s type of A. elegans? 
came from the lower Rio Grande, Texas, and belongs to the 
race occurring from about the ninety-eighth meridian in Texas 
west through northeastern Mexico and New Mexico into south- 
eastern Arizona. For the other race I propose the name 


Arizona elegans occidentalis, new subspecies 


Diagnosis: Similar to A. elegans elegans (Kennicott), but 
with fewer dorsal scale rows, more numerous and narrower 
dorsal blotches, with the lateral spots narrow or indistinct, and 
with relatively shorter tail. 

Type Specimen: United States National Museum No. 54372; 
La Jolla, California; collected by J. C. Thompson, May, 1916. 


1 Contribution from the Zoological Laboratory of the University of 
Michigan. 

? United States National Museum No. 1722, Mexican Boundary Survey, 
Reptiles, p. 18, pl. 13. 


2 University of Michigan 


Description of Type Specimen: Ventral plates, 214; anal 
plate entire; caudal plates, 51, all divided; dorsal scale rows, 
27-25-23-21-19; upper labials, 8 on each side; lower labials, 
13 on the left side and 12 on the right; on each side a single 
preocular and 2 postoculars; temporals, on each side, 2, fol- 
lowed by 4; posterior chin-shields about half as large as the — 
anterior, parallel, and separated by two to three lines of 
small scales; rostral prominent, projecting about half way 
between the internasals; other head shields normal for the 
genus; dorsal scales all smooth, and similar in size and pro- 
portions. 

Total length, 992 mm.; tail length, 136 mm.; tail, therefore, 
0.137 of the total length. Sex, male. 

The general color above (as preserved) is light brownish 

marked by about 72 narrow, ill-defined transverse blotches of 
darker brown. Small lateral alternating spots are faintly 
defined on the anterior half of the body. On the second and 
third, occasionally first and fourth, rows of dorsal scales are 
frequent small black spots, which are rather conspicuous in a 
general view of the specimen. The brown of the upper sur- 
faces changes gradually at about the third or fourth row of 
dorsal scales to the cream color of the lower surfaces. The 
latter are without markings of any kind. The top of the head 
is nearly uniformly hght brown. From the lower postocular 
to the last upper labial is a dark streak, and the labial suture 
under the middle of each eye is dark along its upper portion. 
The under side of the head is unmarked. 
_ Remarks: Specimens from Texas, and from western Arizona 
and California, are referable on sight to their respective sub- 
species, but individuals from the intermediate region are gen- 
erally intermediate in pattern or sealation. A larger series of 
specimens will undoubtedly make possible a closer definition 
of the characters and ranges of these races. Until then a 
specimen must be identified as belonging to the race it most 
nearly resembles in characters and locality. 

The intermediates among the specimens examined are as 
follows: U. S. N. M. 8408 from ‘‘southeastern Arizona’’ has 
the pattern of elegans and the seale rows of occidentalis; U.S. 


Occasional Papers of the Museum of Z oology 3 


N. M. 8002, from Camp Grant, Arizona, has a pattern some- 
what similar to elegans; U. 8. N. M. 14298 from Chihuahua, 
Mexico, has the pattern of elegans and the dorsal scale formula 
of occidentalis ; the specimen from Mesilla Valley, New Mexico, 
is intermediate, as well as four others of indefinite locality 
(A. M. N. H. 4268, 4269, and U.S. N. M. 44904, 48696). 

The two subspecies may be distinguished as follows: 


Seale rows usually 29 or 31; dorsal blotches on body about 55 (40 to 
57) large and squarish, covering about 12 or 13 lateral rows of scales 
and 2 to 3 longitudinal rows, and separated by 1 to 1% scales; lateral 
spots conspicuous and roundish; tail 0.138 to 0.159 of total length. 

A. elegans elegans (Kennicott). 

(From about the 98th meridian in Texas, west through northeastern 
Mexico and New Mexico into southeastern Arizona. 

Seale rows 27, only occasionally 29; dorsal blotches on body about 60 
(54 to 77), narrow, covering about 7 to 10 lateral rows and 1% to 2 
longitudinal rows of scales, and separated. by about 2 scale lengths; 
lateral spots narrow or indistinct; tail 0.100 to 0.148 of total length. 

A. elegans occidentalis, new subspecies. 

(Southeastern Arizona west through southern California and northern 

Lower California. ) 


Seale features may be compared in the accompanying lists 
of specimens. 


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NuMBER 151 JuLy 1, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ArBoR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


SOME PANAMANIAN FROGS! 
By E. R. DuNN 


While Mr. C. B. Duryea and I were collecting for the Mu- 
seum of Comparative Zoology in Panama and Costa Rica in 
the summer of 1923 we were so fortunate as to observe the 
eggs and tadpoles of two species of Hyla, and to collect and 
make observations on the habits of four species of Phyllobates, 
one of them new. The latter was also one of the species col- 
lected by Mr. and Mrs. Gaige for the University of Michigan 
in Panama in the spring of 1923. They worked from the 
Pacific side and we from the Atlantic, but we both collected 
in the cloud forest at about 4,500 feet altitude near the vol- 
eano of Chiriqui and I am under obligations to them for their 
courtesy in allowing me to examine their material and to de- 
seribe the form. I have taken this opportunity to examine all 
available specimens of Phyllobates from Panama and Costa 
Rica. No species of this genus from Central America was 
known to Barbour and Noble when they revised it (Bull. Mus. 
Comp. Zool., LXIIT, 8, 1920). 


1 Contributions from the Department of Zoology, Smith College, No. 
120. 


bo 


University of Michigan 


Unfortunately, Mr. Duryea and I did not discriminate be- 
tween P. talamancae and P. latinasus while in the field, but 
the fact remains that contrary to the general experience oi 
herpetologists all the frog calls we heard were so nearly iden- 
tical that we could not certainly tell which of several forms 
was calling, and this identity of calls included Hyla albomar- 
gyinata, Hyla uranochroa, Phyllobates latinasus and Phyllo- 
bates talamancae at one locality, and the new form from the 
nearby cloud forest. The call of these five forms was a rather 
metallie ‘‘cheep,’’ a single, brief note, repeated at intervals. 
There were unquestionably slight differences, appreciable 
when one watched a calling specimen. Thus Hyla wranochroa 
had a less metallic note than Hyla albomarginata and both 
were louder than the Phyllobates, yet the general impression 
was much the same. 

The two Hylas and two of the Phyllobates were observed 
most carefully at a place called La Loma where we camped 
for a little over a week. This was in rain forest.near the 
trail from Chiriquicito to Boquete, and at an altitude of about 
2.000 feet. Here, in a little brook, we found four sorts of tad- 
poles. Two of these we traced to two Hylas which we had 
previously caught; the other two later proved to be Phyllo- 
bates. Two lots of eggs were found, and as the Phyllobates 
males carried their tadpoles to the stream, it seems fair to 
assume that the eggs were those of the two Hylids. The four 
tadpoles from La Loma were all traced to their respective 
adults by complete series of transforming young. 


Hyla uranochroa Cope 


Eggs assumed to be those of this species were found at- 
tached to a leaf overhanging the stream. The eggs when 
found contained well developed tadpoles. This form of egg- 
_ laying has been reported for Agalychnis and for Phyllome- 
dusa. Hyla uranochroa has much the appearance of an Aga- 
lychnis, as it is uniform green, with very prominent red eyes. 
The technical characters are those of Hyla. The species was 
never seen on the ground or in the water, and the calling indi- 
viduals were always on leaves; even the transforming speci- 


Occasional Papers of the Museum of Zoology 3 


mens were caught in such situations. It would seem that Hyla 
uranochroa might well be an approach towards the two more 
specialized genera in eye color and in habits, while in form of 
pupil and character of hand still a technical Hyla. 

The tadpoles of this species were the first to be assigned 
correctly to their adults, on account of the red eyes, which 
were a prominent feature of both stages. They were very 
common in the stream and possessed well developed power of 
suction. We often lifted them from the water clinging to our 
hands in this way. They were occasionally observed clinging 
to rocks or to the surface film. 

Diagnostic characters of mature tadpole: Spiracle sinistral, 
anus dextral, eyes visible from ventral surface, upper fin crest 
not extending to anus, spiracle halfway between snout and 
base of hind limb, labial teeth 2/3, uniform brown, eyes red, 
greatest length 45 mm., smallest seen 16 mm. 

Description of mature tadpole: Length of body contained 
three times in total length; width of body 1.75 times in its 
own length; nostril nearer to eye than to snout; eye dorso- 
lateral, visible from ventral surface, equidistant from snout 
and from spiracle; distance between nostrils — interorbital 
width = width of mouth; spiracle sinistral, equidistant from 
snout and from base of hind legs; anus dextral; depth of the 
muscular portion of tail at base 1.2 in greatest depth of tail. 

Upper labium with two equal uninterrupted rows of teeth; 
lower labium with three continuous rows, inner two equal, 
outer about 14 as long as median; a complete lip around the 
mouth. 

General color, uniform brown; crests of tail unpigmented 
save for a few spots on the dorsal portion. Total length 43 
mm., tail 30 mm. 


Hyla albomarginata Spix 


A foamy mass of eggs was found under a rock in the small 
stream. As this was the situation from which Hylt albomar- 
ginata was ealling, it is quite possible, by association and by 
elimination of the other three species whose tadpoles inhabited 
the stream, that these were the eggs of this animal. 


4 University of Michigan 


The dark brownish green adults were all taken under rocks 
in the water, whence they were calling. One transforming 
specimen was on a leaf. This was bright green with tiny dark 
dots, and a white line from the eye over the tympanum. 

Diagnostic characters of mature tadpole: Spiracle sinistral, 
anus dextral, eye not visible from ventral surface, upper fin 
crest not extending beyond hind curve of body, distance from 
spiracle to base of hind limb contained 1.5 times in its dis- 
tance from the snout, labial teeth (6-7/8), brown above, light 
cross bars on dorsum of tail, crests unpigmented, greatest 
length of tadpole 60 mm. 

Description of mature tadpole: Length of body contained 
2.9 times in total length; width of body 1.5 in its own length; 
nostril nearer to eye than to snout; eye dorsal, nearer to snout 
than to spiracle; distance between nostrils greater than inter- 
orbital width, less than width of mouth; spiracle sinistral, its 
distance from the base of the hind limb contained 1.5 times 
in its distance from the snout; anus dextral; depth of the mus- 
eular part of tail at base contained 1.5 times in greatest 
depth of tail. Upper labium with 7 equal rows of teeth, the 
outer three somewhat fragmented, the innermost narrowly 
broken in the middle; lower labium with eight rows of teeth, 
of nearly equal length, the outer four somewhat fragmented ; 
a complete circlet of papillae around mouth; in repose this 
contracts into a triangle, apex forwards, lateral angles curved 
in and back. The general color was brown, with three lighter 
reddish brown cross bars on the muscular part of the tail, the 
erests unpigmented, and the belly pale; total length 53 mm., 
head and body 18 mm., tail 35 mm. 

These tadpoles were most frequently seen in numbers cling- 
ing to the rocks over which a swift current of water was 
streaming. They had by far the best developed sucking disks 
of the four forms in the stream. 


Phyllobates 


Barbour and I (Proce. Biol. Soe. Washington, 34, p. 159, 
1921) redeseribed Phyllobates talamancae (Cope) from two 
specimens from Santa Cecilia, Costa Rica, and described 


Occasional Papers of the Museum of Zoology 5 


Phyllobates beatriciae as new with a single specimen from 
Zent, Costa Rica. On the basis of more extensive material and 
re-examination of these specimens the opinions we then ex- 
pressed require modification. 

Phyllobates beatriciae, of which five additional specimens 
were taken at Almirante, Panama, and four in the Talamanca 
Valley, Costa Rica, by Duryea and myself, seems to be iden- 
tical with the frog described as Dendrobates lugubris by 
Schmidt (Denksch. Acad. Wien, 14, 1858, p. 250, pl. 2, fig. 
14). In the specimen we described the yellow dorso-lateral 
lines had disappeared, and so had the line from the arm to 
under the eye. These are quite apparent in the series at hand. 
Schmidt’s specimens are well described and figured, and while 
his are said to have come from the cloud forest at 5,000-7,000 
feet, and ours all came from low rain forest, there seems to be 
no doubt that the two are identical and the animal must be 
known as Phyllobates lugubris (Schmidt). We did not take 
it at La Loma nor in the cloud forest. It was observed to 
have the tadpole-carrying habit. Its very different coloration, 
dense black with narrow yellow lines, precludes any confusion 
with the other forms. 

The redeseription of Phyllobates talamancae by Barbour 
and myself was based on a male of this species and a female 
of what seems to be Phyllobates latinasus (Cope). This speci- 
men of latinasus was assumed to be a female at the time of 
describing. The male of talamancae has a black throat and 
this specimen was carrying tadpoles; the very similar speci- 
men of datinasus, taken in the same stream, with a light throat, 
and without tadpoles, was assumed to be the female, an as- 
sumption which dissection has shown to be erroneous. 

The two species were found in both Costa Rica and Panama 
last summer, and were supposed to be the same. Two very 
different sorts of tadpoles at La Loma, which both seemed to 
us in the field to turn into the same frog, caused a critical 
examination of all the available material upon our return. 
Specimens of Phyllobates latinasus have been seen from Santa 
Cecilia, Costa Rica; Almirante and La Loma in western Pan- 
ama; Cerro Azul near the Canal Zone (U.S. N. M., 54174-5) ; 


6 University of Michigan 


Cana (U.S. N. M., 54231, 63005) and Rio Esnape in extreme 
eastern Panama. The last two lots are so near the type loeal- 
ity, the Truando region of Colombia, as to be virtually topo- 
types. 

P. talamancae has been seen from Santa Cecilia and from 
Suretka, Costa Rica, and from La Loma in Panama. It was 
described from Old Harbor on the coast between Limon and 
Almirante. 

The most obvious difference between the two is the marking 
of the sides. In both species the dorsal region is gray and the 
sides are an intense black. There is an ill-defined light line 
where the gray and black meet. In both species a white streak 
starts from the groin and passes obliquely forward and up- 
ward toward the upper eyelid. In latinasus it dies out about 
halfway between leg and arm, but in talamancae it reaches the 
eye, and rather supersedes the ill-defined line which separates 
the black from the gray. The result is that talamancae ap- 
parently has a single white streak on the side and that latina- 
sus has one and a half. 

The marking of the thigh in talamancae is a hooked or 
‘‘anvil-shaped’’ affair, owing to the dark line along the dorsal 
surface of the thigh running into a black area in the knee-pit 
which extends along the posterior aspect of the thigh. In lati- 
nasus this latter is absent and there is merely a dark streak 
on the dorsal surface. 

The legs of latinasus are barred, while the legs of tala- 
mancae are not barred. 

There are various differences in the length of toes and size 
of disks. These differences are essentially that the inner and 
outer toes of talamancae are reduced. Thus the tip of toe I 
reaches the penultimate joint of II in latinasus but not in tala- 
mancae and beyond it in latinasus. Two phalanges of IV are 
in latinasus but only the antepenultimate in talamancae. The 
tip of IIIT reaches the antepenultimate joint of IV in tala- 
mancae and beyond it in latinasus. Two phalanges of IV are 
beyond the tip of V in latinasus and 214 in talamancae. The 
tip of V reaches the penultimate joint of III in talamancae 
and beyond it in Jatinasus. The disk of V is equal to half the 


Occasional Papers of the Museum of Zoology r’ 


disk of IV in talamancae, but in latinasus they are the same 
size. 

There is marked sexual dimorphism in talamancae. The 
throat of the male is black and the third finger of the male is 
so swollen that the disk is no larger than the rest of the finger. 
These characters are not apparent in latinasus. 

Finally, while Jatinasus has the normal tadpole of the genus, 
with labial teeth 2/3, as described for subpunctatus, sylvatica, 
and trinitatis, the tadpole of talamancae has no labial teeth, 
but a highly developed labial disk, like that of Microhyla 
achatina and Megalophrys montana and entirely different 
from anything hitherto described from America. 

Phyllobates kingsburyi Boulenger, a large species with gray- 
ish mottling on throat and chest, seems to occur east of the 
Canal Zone. Specimens considered to be this form have been 
seen from Rio Calobre about south of the Gulf of San Blas 
(U. S. N. M., 53737-8), and from Cana (U. S. N. M., 50177, 
50197-50200). 

In the cloud forest above the Chiriqui lagoon, on the slopes 
of Chiriqui and Horqueta, and in the drier forest down as far 
as Boquete lives another species allied to talamancae and hav- 
ing the same sort of tadpole. It may be called 


Phyllobates nubicola, new species 


Diagnosis: Closely related to P. talamancae (Cope) from 
which it may be distinguished by the following characters: 
the black line on the dorsal surface of the thigh is not hooked, 
the thigh is red in life and the belly yellow, the tip of toe 
V reaches past penultimate joint of toe III, two phalanges of 
toe IV reach beyond the tip of toe V, the tip of toe I reaches 
the penultimate joint of toe IT. 

From P. latinasus it may be distinguished by the white line 
from the groin reaching the eye, and by the disk of toe V being 
half the size of the disk of toe IV, as well as by the black 
throat and swollen third finger of the male, in which charac- 
ters it agrees with talamancae. 

Type specimen: Cat. No. 58292, Museum of Zoology, Univer- 
sity of Michigan. Adult male, collected in rain forest above 


8 University of Michigan 


Boquete on the trail to Chiriqui Grande, 4,500 feet, by F. M. 
Gaige, May 15, 1923. 

Description of type specimen: Snout moderate, as long as 
orbit ; loreal region vertical, somewhat concave; nostril nearer 
tip of snout than eye; interorbital space much broader than 
upper eyelid; tympanum indistinct, 1/3 the size of eye; disks 
well developed, smaller than tympanum; third finger swollen, 
as wide as disk; disk of toe I half as large as that of toe II; 


yj Ry, 


Right forefoot of male of Phyllobates nubicola, ventral surface, x74. 


disk of toe V half as large as that of toe IV; tip of toe I 
reaches penultimate joint of toe II; tip of toe II reaches ante- 
penultimate joint of toe III; tip of toe III reaches antepen- 
ultimate joint of toe IV; two phalanges of toe IV beyond tip 
of toe V; tip of toe V reaches past penultimate joint of toe 
III; two small metatarsal tubercles; a tubercle in the middle 
of the tarsus; tibiotarsal articulation reaching eye; skin 
smooth. Head and body black above and on sides; a light line 
from above eye to groin; a light line under eye above insertion 
of arm to groin; chin and throat black, belly mottled black and 
white; a dark line on anterior edge of thigh; a dark line on 
dorsal surface of thigh; a dark triangle, apex at vent, corners 
along thigh; light portions of thigh red, legs and arms uni- 
form dusky; snout to vent 21 mm.; head width 7 mm.; head 
length 7 mm.; arm 13 mm.; leg 28 mm. 

Variations: Females are white below. Many males have 
only the throat dark. Many specimens have the lower light line 
continuous around upper lip. Specimens seen: 30, as follows: 
Cloud forest above Gutierrez, 6,000 feet on trail from Chiri- 


Occasional Papers of the Museum of Zoology 9 


quicito to Boquete, Atlantic slope, 6; Pacific slope, same trail, 
under stones and debris along edge of stream 2 miles above 
Boquete, 3,900 feet, 11 (U. Mich., 58286) ; same trail, wet for- 
est 4,500-5,500 feet, 8 (U. Mich., 58289, 58291) ; rain forest 
by last creek on trail on Pacific slope, 4,500 feet, 5. 


Key to Central American forms of Phyllobates 
A. Dorso-lateral line yellow, throat of both sexes black, belly black 
with fine yellow lineg................ P. lugubris (Schmidt), (Costa Rica 
and western Panama). 
AA. Dorso-lateral line white. 
B. Throat and chest of both sexes mottled with gray, disk of toe 
disk s0 f. GOS? Woe eens: P. kingsburyi Boulenger (eastern 
Panama to Ecuador). 
BB. Throat and chest of female always immaculate, disk of toe I= % 
disk of toe II. 
C. White line from groin not reaching eye, disk of toe V = disk of 
toe IV, throat of male white............... P. latinasus (Cope) (Costa 
Rica to Colombia). 
CC. White line from groin reaching eye, disk of toe V—=% disk of 
toe IV, throat of male black. 
D. Anvil-shaped black marking on thigh, thigh not red in life, tip 
of toe V not reaching penultimate joint of toe IV.....P. talamancae 
(Cope) (Costa Rica, western Panama). 
DD. Linear black marking on thigh, thigh red in life, tip of toe V 
reaching penultimate joint of toe IV.........0.. P. nubicola Dunn 
(western Panama at high altitudes). 


Save for P. kingsburyi we saw all these carrying their 
tiny tadpoles. Tadpoles of latinasus, talamancae, and nubi- 
cola are at hand. Similar habits have been described for 
trinitatis and for swbpunctatus. Apparently the male carries 
the very young tadpoles to the stream and leaves them there. 
There is no evidence that the male of Phyllobates gets the tad- 
poles from the water. The stream at La Loma contained all 
stages from the tiny young ones 10-12 mm. long to the trans- 
forming specimens. Males carrying tadpoles were put into 
water and the tadpoles left the parent and swam away. Noth- 
ing was seen of the eggs or the mating. The tadpoles of latt- 
nasus are similar in all respects to the described tadpoles of 
sylvatica, trinitatis and subpunctatus. The tadpoles of tala- 
mancae and of nubicola are very different. 


10 University of Michigan 


Diagnosis of the mature tadpole of P. latinasus: Spiracle 
sinistral, anus dextral, eye invisible from the ventral surface, 
upper fin crest extending to end of body, distance from spir- 
acle to base of hind limb contained 1.2 times in its distance 
from the snout, labial teeth 2/3, light grayish brown, fine dots 
on tail and on crests, greatest length 36 mm. 

Description of mature tadpole of P. latinasus: Length of 
body contained 2.9 times in total length; width of body 1.4 in 
its own length; nostril equidistant between eye and snout; 
eye dorsal, not visible from below, nearer to snout than to 
spiracle; distance between nostrils —interorbital width= 
width of mouth; spiracle sinistral, its distance from the base 
of the hind legs 1.2 times in its distance from the snout; anus 
dextral; depth of the muscular portion of the tail contained 
twice in greatest depth of tail. Upper labium with two series 
of teeth equal in length, inner divided in the middle for a dis- 
tance equal to 14 the length of either half; lower labium with 
three equal series of teeth; papillate border of mouth broadly 
absent anteriorly, upper row of teeth forming border; light 
grayish brown above, darker spots on base of tail and on 
crests. Total length 32 mm., head and body 11 mm, tail 
21 mm. 

Diagnosis of mature tadpole of P. talamancae: Spiracle sin- 
istral, anus dextral, eye invisible from the ventral surface, 
upper fin crest well developed only on distal 2/3 of tail, spir- 
acle equidistant between snout and base of hind limb, no labial 
teeth, brownish with a black streak on upper edge of muscular 
part of tail, greatest length 33 mm. 

Description of mature tadpole of P. talamancae: Length of 
body contained 3.4 times in total length; width of body 1.3 in 
its own length; nostril equidistant between eye and snout; eye 
dorsal, not visible from below, nearer to snout than to spir- 
acle; distance between nostrils less than interorbital width, 
less than width of mouth; spiracle sinistral, equidistant be- 
tween snout and base of hind legs; anus dextral; depth of 
muscular portion of tail at base contained 11, times in great- 
est depth of tail; no labial teeth; a broad labial disk, the pos- 
terior portion of which is twice as deep as the anterior, papil- 


Occasional Papers of the Museum of Zoology 11 


late throughout; brown, posterior half of tail with black line 
on upper edge of muscle, some mottlings on other parts of 
muscle and a few dots on erests; total 31 mm., head and body 
9, tail 22. Many of the tadpoles are lighter in color than the 
described one but in all the dark line on the tail can be made 
out. 

These two sorts of tadpoles were common in the stream at 
La Loma. Those of latinasus were similar to Rana tadpoles in 
appearance and in actions. The tadpoles of talamancae seemed 
to remain in the depest parts of the pools and were very active 
and darter-like in their movements. 

Three tadpoles from near Boquete: ‘‘under low stones in 
washed out cavity at edge of small stream, 4,100 feet,’’ col- 
lected by the Gaiges, February 26, 1923, are much like the 
tadpoles of talamancae, but with quite obvious differences. 
They are probably the larvae of nubicola. 

Diagnosis of mature tadpole of P. nubicola: Spiracie sinis- 
tral, anus dextral, eye invisible from the ventral surface, 
upper fin crest well developed only on distal 2/3 of tail, dis- 
tance from spiracle to base of hind limb contained 1.2 times 
in its distance from the snout, labial teeth 1/0, uniform 
brown, largest of the three 32 mm. long. 

Description of mature tadpole of P. nubicola: Length of 
body contained 3.2 times in total length; nostril nearer snout 
than to eye; eye dorsal, not visible from below; eye nearer 
snout than to spiracle; distance between nostrils = interor- 
bital width, less than width of mouth; spiracle sinistral, its 
distance from the base of the hind limb contained 1.2 times 
in its distance from the snout; anus dextral; depth of mus- 
cular portion of tail at base is the greatest depth of tail; labial 
teeth 1/0; the row of teeth not as long as width of beak; a few 
teeth between tooth row and beak perhaps represent a rudi- 
mentary inner upper row; a broad labial disk, posterior part 
twice as deep as anterior, papillate throughout; brown, prac- 
tically uniform, crests pale; total length 32 mm., head and 
body 10 mm., tail 22 mm. 

In both adult and tadpole nubicola is intermediate between 
latinasus and talamancae. 


12 University of Michigan 


Matters of considerable general biological interest are sug- 
gested by the forms described above. A Hyla with the habits 
but not the structure of an Agalychnis; Phyllobates whose 
adults are so similar as to have been considered conspecific by 
three observers but whose tadpoles differ in the most extra- 
ordinary fashion; these support the precedence in time of 
function over form, and the possibility of apparent reversal of 
the biogenetic law, owing to different larval habits and similar 
adult habits. 

The case of Megalophrys seems much like that of Phyllo- 
bates. Adult and larva live in two different worlds, and while 
the general environment and habits of most species are more 
alike as larvae than as adults, the reverse may be, and in these 
eases undoubtedly is, true. 

The much more profound changes which attend metamor- 
phosis in frogs than in salamanders lead one to marvel why 
neoteny is unknown in the former while fairly common in the 
latter. 


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14 


University of Michigan 


PLATE I 


Tadpole of Hyla uranochroa, side view, x2. 
Mouth of tadpole of Hyla uranochroa, x6. 
Tadpole of Hyla albomarginata, side view, x2. 


Mouth of tadpole of Hyla albomarginata, x6. 


PANAMANIAN F'Rocs 


PLATE 


16 


One 


University of Michigan 


PLATE II 
Tadpole of Phyllobates talamancae, side view, x2%. 
Mouth of tadpole of Phyllobates talamancae, x7. 


Tadpole of Phyllobates nubicola, side view, x2%. 


. Mouth of tadpole of Phyllobates nubicola, x7. 


Tadpole of Phyllobates latinasus, side view, x24. 


Mouth of tadpole of Phyllobates latinasus, x7. 


PANAMANIAN F'RoGs PLATE II 


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NUMBER 152 Avucust 12, 1924 


OCCASIONAL PAPERS OF THE MUSEUM OF 
ZOOLOGY 


UNIVERSITY OF MICHIGAN 


ANN ARBOR, MICHIGAN PUBLISHED BY THE UNIVERSITY 


LAND AND FRESHWATER MOLLUSCS OF THE 
DUTCH LEEWARD ISLANDS 


By H. Burrincton BAKER 


INTRODUCTION 


In 1920, while on the University of Michigan-Williiamson 
Expedition to Venezuela, a small collection of land shells was 
obtained from the Schaarlo, back of Willemstad, Curacao.* 
In 1922, the University of Michigan Museum of Zodlogy sent 
me back to the Dutch Leeward Islands to collect molluses, 
reptiles, amphibians and ants. With the assistance of an addi- 
tional grant from the Zodlogical Laboratory of the University 
of Pennsylvania, for the collection of grasshopper testes as 
cytological material, I was able to spend the summer, from 
June 11th to September 17th, 1922, in a study of all five of 
the islands of the group. The reptiles collected have been 
listed already, in a paper by Dr. A. G. Ruthven.? 

From June 11th to 28th, Overzijde (Otrabanda), Curacao, 
was the base for collecting trips. From June 29th to July 


11923; Occ. Papers, Mus. Zool. Univ. Mich., no. 137; pp. 1-7, pl. I. 
21923; Occ. Papers, Mus. Zool. Univ. Mich., no. 143. 


2 University of Michigan 


12th, I was guest of Mr. Richard Muskus at his Landhuis 
Knip, near the northern end of the same island. On July 19th, 
the schooner ‘‘Albertina’’ took me to Oranjestad (Plaja 
Aruba), from which town a study was made of the island of 
Aruba, until August 10th, when I sailed, on the packet 
schooner ‘‘ Ligia,’’ back to Willemstad (Punta), Curacao, and 
then to Kralendijk (Plaja Bonaire), Bonaire. From August 
13th to September Ist, collections were made on this island, 
with a visit to Klein-Bonaire (Bonaire Chikitoe) on August 
24th. By arrangement, the sloop ‘‘America’’ put me ashore 
for two hours, on September Ist, at the island of Klein- 
Curacao (Curacao Chikitoe). September 2nd to 17th were 
spent at Overzijde, Curacao, from where trips were made to 
Sint Willebrordus and New Port, on the same island. 

A note in regard to the locality names may not be out of 
place. The language of the Dutch Leeward Islands is 
Papiamento, which seems to have started as a dialect of 
Portuguese, but which has acquired words from all of the 
languages spoken in the West Indies. Written Papiamento 
was invented by Dutch orthographers, with the result that 
the combinations of letters, used to express certain sounds, 
are very different from those in the Romance languages. The 
larger towns and the more conspicuous topographic features 
have Dutch names, but the official language is infrequently 
heard in the islands, and the Papiamento synonyms are much 
more commonly used. The former are preferred throughout 
this paper, but the latter are occasionally added in paren- 
theses. The spellings used here are taken from the Dutch 
Government 1/20000 topographic maps, but, even in these, 
variations occur. 

My thanks are due the Government officials of the islands, 
especially the Procureur-General and the Government seecre- 
taries of Curacao and the Subgovernors of Aruba and Bonaire, 
and also to the United States Consul, Mr. B. S. Rairden, for 
assistance in many ways. In addition, I found all of the 
people of the islands extremely hospitable and always willing 
to direct me to favorable localities for study. In particular, 


Occasional Papers of the Musewm of Zoology 3 


I wish to express my indebtedness to Mr. Richard Muskus, of 
Campo Knip, Curacao, whose hospitality I enjoyed for two 
weeks, to Mr. Gravenhorst, of Kralendijk, Bonaire, who helped 
me to find quarters on that island, and to Mr. and Mrs. de 
Veer, of Oranjestad, Aruba. I also wish to thank the firm of 
S. E. L. Maduro and Sons for their many courtesies, and that 
of John Godden and Co., for permission to visit the Tafelberg 
of Santa Barbara. 

I am deeply indebted to Ir. G. J. H. Molengraaff, M.I., for 
the meteorological data which are collated in Tables I, II and 
III, and to Dr. N. L. Britton, who, through the kind mediation 
of Mr. J. M. Fogg, identified some specimens of characteristic 
plants. The identification and comparison of the molluses col- 
lected were made possible by the library and collections of the 
Academy of Natural Sciences of Philadelphia, where Dr. H. 
A. Pilsbry and Mr. E. G. Vanatta were, as always, very help- 
ful. Drawings, photographs, and preparations of radulae, ete., 
were made at the Zodlogical Laboratory of the University of 
Pennsylvania. 


ENVIRONMENT 


The Dutch West Indies, or the Netherlands Colony of 
Curacao, consist of islands in the northern Lesser Antilles, and 
the ones studied in the Leeward Group. The island of 
Curacao, from which the entire colony takes its name, is the 
largest of the latter, and lies in the Caribbean Sea, 47 miles 
north on the 69th west meridian from the coast of Venezuela, 
and just north of 12° north latitude. The island of Aruba 
is about 60 miles to the northwest; it lies on the 70th west 
meridian, approximately -20 miles north of the Paraguana 
Peninsula, and around 12° 30’ north latitude. Bonaire and 
the closely associated island of Klein-Bonaire are about 30 
miles east of Curacao, just northwest of 12° north latitude and 
68° west longitude. Klein-Curacao is a small coral island 
about 8 miles southeast of the southeastern end of Curacao, 
at 12° north latitude and 68° 39.5’ west longitude. 

The ocean bottom has not been mapped thoroughly in all 
parts adjacent to the islands, but it seems quite certain that 


4 University of Michigan 


738 fathoms of water lie between Curacao and the mainland. 
Such a depth of water would mean that a constant negative 
movement of the strand line would connect the Lesser Antilles, 
and even the Greater Antilles, to the South American main- 
land, before the 40 mile strait between Curacao and Venezuela 
would be drained. On the other hand, the passage between 
Aruba and the Paraguana Peninsula is less than 40 fathoms 
deep in the shallowest place. The United States Hydro- 
graphie Office chart, number 964, shows one sounding of 830 
fathoms between Curacao and Aruba, but the depths between 
Curacao and Bonaire, and between the last island and the 
groups to the east, are not mapped. Nevertheless, it seems 
probable that the isolation of Curacao, Klein-Curacao, Bonaire 
and Klein-Bonaire is at least directly comparable to that of 
the northern Lesser Antilles. As will be discussed later, this 
geographic separation is intensified by the ecological differ- 
ences between all of the Dutch Leeward Islands and most parts 
of the South American mainland. 

Ir. G. J. H. Molengraaff, M.I., Chief of the Weather 
Bureau of Curacao, very generously sent me the meteorological 
data which are compiled into the following tables. As will be 
seen from the first of these, the climate of the Dutch Leeward 
Islands is influenced by an almost constant, east trade wind 
of considerable strength. As will be discussed later, this pro- 
duces a marked difference between the eastern and western 
slopes on the islands. All of the records given are from points 
on the western side of the islands, with the exception of those 
from Rincon, Bonaire, which is also partially protected from 
the east winds. 

The usual ocean currents follow quite closely the direction 
of the trade winds, but are deflected somewhat to the north- 
ward by the South American coast line and those of the 
islands. For instance, the trip from Willemstad to Oranjestad 
was accomplished in about 12 hours, although the schooner’s 
sails were flapping most of the time. On the other hand, the 
return trip, which was said to be a fast one, required 34 hours. 
However, this general trend is occasionally reversed. ‘‘South 


Occasional Papers of the Museum of Zoology 5 


Table I. Winds at Fort Amsterdam, Willemstad, Curacao 
from August, 1910, to December, 1921, inclusive 
Wind Velocity 


Wind Direction in Percentages Beaufort Scale (0-12) 
8:00 2:00 6:00 


N. N.E. E. S.E. S. S.W. W. N.W. A.M. P.M. P.M. 
J AMUATY eens. 0.2 14.8 84.0 0.7 0.0 0.1 02 0.0 2 2 


September ...... 0.1 62 87.8 5.0 04 0.2 0.2 0.0 
October .....w...... Ose erSto moO e Weal a0 ales Ole. OSE 
November ...... Ose lo 4aere:0) (6:00 120) lp 0227 20.0 
December .......... (O40), aBRBy cB er SP a Kone COM) Ces Oe 
Mean Annual 0.2 10.3 85.7 29 04 04 0.1 0.0 


February ......... 06 16.0, 642° 1.9 G:0 0.0 00 O1 3 3 
March 13.9 84.2 1.9 0.0 0.0 0.0 0.0 3 3 
April 98 869 13 0.2 1.2 0.0 0.0 3 3 
May b=... , 8.8 88.0 3.2 0.0 0.0 0.0 0.0 3 3 
June ........ : 6.7 92.6 0.7 0.0 0.0 0.0 0.0 4 3 
Sul? ee as l ta ote. 1 Os OF OL G0: 0.0 3 3 
August 74 88.2 3.4 01 05 0.1 0.0 3 3 
3 3 
2 2 
2 2 
3 3 
3 3 


Ww bd dD Ww Ww Ow Co Ww OO DO DO 


of Curacao the surface current is generally to the westward, 
but an easterly sub-current exists and this is of such volume 
that it is liable entirely to overcome the surface set.’’* 

The temperatures (Table II) are remarkably uniform, with - 
a total extreme fluctuation of 23 degrees Fahrenheit, during 
the period measured. The mean annual temperature (81° F.) 
places the islands in Calvert’s zone II. It is also noteworthy 
that the mean daily variation (8° F.), although small, is twice 
as great as the fluctuation in the mean monthly temperature 
ca” Ey). 

The rainfall (Table III) is as variable as the temperature is 
uniform. The mean annual rainfall of the four localities 
measured (17 to 22 inches) immediately establishes the islands 
as tropical semi-deserts. It will be noted that a distinct rainy 
season occurs in the months October to January, although the 


3 Observations missing between Nov. 2nd and 7th, 1918. The table of 
wind directions is reduced from one containing intermediate points; most 
of the N.E. and S.E. winds listed were actually observed as E.N.E. and 
E.S.E. winds, respectively. 

4 United States Hydrographic Office chart, number 1290. 

51908; Proc. Acad. Nat. Sci. Philadelphia; Plate xxvi. 


6 University of Michigan 


Table II. Temperatures at Fort Amsterdam, Willemstad, Curacao 
from January, 1903, to June, 1921, inclusive’ 


Mean Monthly Mean Daily Range Extremes 

Cent. Fahr. Centigrade Fahr. Centigrade Fahr. 
January ............. 25.9 79 23.6-28.2 74-83 21.0-30.5 70-87 
February ............ 25.9 79 23.7-28.2 75-83 21.0-30.5 70-87 


79 23.7-28.7 75-84  20.5-31.0 69-88 
80 24.5-28.7 76-84 21.0-31.5 70-89 
82 25.4-30.1 78-86 23.0-32.8 73-91 
82 25.6-29.9 78-86  22.4-32.0 72-90 
82 25.3-29.9 78-86  22.0-32.2 72-90 


PAUSE ences 26.7 80 25.7-30.3 78-87 21.0-32.5 70-90 
September ......... 28.3 83 25.9-30.8 79-87 20.0-33.0 68-91 
Getoper, =... % 28.0 82 25.6-30.4 78-87 21.0-33.0 70-91 
November ........... 27.5 81 25.1-29.8 77-86 21.0-32.0 70-90 
December ............ 26.5 80 24.2-28.8 76-84  21.5-32.3 71-90 
ATINUAL, Leese 27.0 81 24.9-29.5 77-85 20.0-33.0 68-91 


usual rainy period of the higher West Indian islands and the 
Venezuelan mainland comes between June and November. 
But extreme variation is the most conspicuous feature; less 
rain may fall in one of the wet months than the mean precipi- 
tation of the driest month. The rainfall of a single month 
may be much greater than that of an extremely dry year; in 
fact, it may be practically as great as the mean annual precipi- 
tation. In addition, my own experience, and conversation 
with the inhabitants, lead me to believe that the greater pro- 
portion of the downfall comes in a few torrential deluges. 
Attention also should be called to the fact that my study of 
the islands was made during the dry season, but that the pre- 
ceding year and the one of my visit (1921 and 1922) were wet 
years (875 and 810 mm. rainfall, respectively, at Cas Chikitoe, 
near Willemstad, Curacao). 

All of the recorded data come from near the towns. From 
the vegetation, and the rainfall while I was on the islands, I 
suspect that the vicinity of the higher hills receives a consider- 
ably greater amount of precipitation, especially during the 


6 Not recorded from Feb. 11-17, 1905; Nov. 3-7, 1918; Sept. 1-23, 
1904; Oct. 5-31, 1904; June and Sept., 1906; July and Sept. to Dec., 
1919; Jan., 1920. 


Occasional Papers of the Museum of Zoology 7 


drier months. In fact, it seems probable that the differences 
between the stations recorded are quite largely due to such 
local factors as the height of the hills in their vicinity. How- 
ever, none of the ‘‘seroes’’ is high or large enough to produce 
anything like a rain forest. Sint Christoffelberg (1,229 feet), 
on its western side near the summit, is the only place where 
bromeliads and other epiphytes are at all conspicuous. 


Table III. Rainfall in Millimeters? 


Fort Amster- Oranjestad, Kralendijk, Rincon, 
dam, Curacao Aruba Bonaire Bonaire 
Mean Extremes Mean Extremes Mean Extremes Mean Extremes 
January ...... 54 899-115 50 = 0-151 50 3-111 63 0-167 
February .. 30 1-102 14 0-64 32 0-95 32 1-79 


March  ......... Py sel | 15 0-56 24 0-74 18 Tr.—50 
J. yal le Sea 24 0-173 23 ~=0-168 24 0-104 28 Tr—185 
Matye 222 ll = 0-54 8 0-51 13 0-48 21 Tr—136 
ere ht 19 1-72 13 (0-54 10 0-30 14 Tr-53 
Jjil) 30 2-89 21 2-45 27 4—62 27 Tr—60 


August ........ 38 3-142 ao atl—139 28 3-88 29 Tr—98 
September 27 4-89 24 2-59 15 1-38 23 0-70 
October ...... 99 1-472 74 6-252 100° Tr—494 88 8-342 
November 118 12-325 96 4358 114 31-241 123 35-214 
December 83 10-262 65 11-228 80 21-265 79 25-269 
Annual... 561 435 86-941 516 164-982 545 133-883 


The data sent me by Ir. Molengraaff show a rather high and 
constant humidity, during the daytime, at Fort Amsterdam, 
Willemstad, Curacao. The mean annual (August, 1910, to 
December, 1920) and the variations in the mean monthly rela- 
tive humidity are: at 8 A. M., 74 and 73~—75, respectively ; at 
2 P. M., 71 and 68-72; at 6 P. M., 73 and 71-74 per cent. 
Although I have no data to present, I believe that dew is a 
comparatively rare and inconspicuous phenomenon on the 


7 The records from Fort Amsterdam, Willemstad, Curacao, cover a 
period of from 28 to 29 years, dependent on the month (Jan. and Feb., 
1895-1923; Mar. to May, 1895-1922; June to Sept., 1894-1922; Oct. to 
Dec., 1894-1921). From Oranjestad, Aruba, they cover a period of 16 
years (1901-1916). From the two localities on Bonaire, they cover a 
period of 12 years (1905-1916). 


a University of Michigan 


highlands and along the western sides of the islands, at least 
during the dry months. On the other hand, along the exces- 
sively arid eastern shores and northern and southern tips of 
the islands, a rather heavy dew formation seems to be the 
usual thing. 


LAND HABITATS 


Sometime after the Miocene,® the Dutch Leeward Islands 
must have been almost entirely submerged; so that they were 
largely covered with a thick coral formation. Three or four 
prominent beach-lines, around the hills (fig. ii-5), were 
formed at different stages of the emergence, and the limestone 
was eroded away from the central portions, so that the older 
rocks were extensively exposed. Recently, another period of 
partial submergence must have taken place, as the peculiar 
shape of such bays as the Schottegat (Plate I, 5-R) and 
Spaansch Water (2—-E) on Curacao, certainly indicate that 
these are sunken valleys. As the central dome was eroded 
away, the stream channels formed narrow outlets through the 
tilted limestone at the edges, and most of the present valleys 
have a similar shape to these submerged ones. 

Fundamentally, on the basis of their igneous intrusions, the 
Dutch Leeward Islands have been considered a portion of a 
complex which includes the Goajira Peninsula and the Sierra 
Nevada de Santa Marta (perhaps also the Paraguana Penin- 
sula) of the South American mainland.’ Superficially, each 
island is a canoe-anticline, the long axis of which extends 
approximately northwest southeast. The central portion of 
each island is composed largely of highly metamorphosed and 
crumpled, Cretaceous conglomerate-schists, and of igneous 


8 Compare J. Lorié; 1887-9; Samml. Geol. Reichsmus. Leiden, 2 ser., 
Bd. I, pp. 111-149; and K. Martin; 1887, 1888; Bericht iiber eine Reise 
nach Niederlindisch West-Indien, I and II. ~The latter contains a de- 
tailed discussion of the geology of the three main islands, with rather 
inelaborate maps of the formations. 

9 W. Sievers; 1896; Petermanns Mitteilungen, Vol. XLII, p. 129, and 
pl. X. 


Occasional Papers of the Museum of Zoology 9 


rocks, such as diabase, quartz-diorite (northern Aruba) and 
mica-porphyrite (northern Bonaire). These older rocks may 
resist erosion so as to form the highest hills in the islands (figs. 
ili-6, and vi-14), or they may be peneplainized (especially 
the diabase) to rolling plains (fig. vii-17), inside of the lime- 
stone ramparts (fig. iii-5) along the seashore. Most of the soil 
of the islands is derived from these more ancient rocks, but the 
rapid erosion prevents the retention of much residue, except 
on the more nearly level portions. As a result, almost all of 
the agricultural development (hofjes), except the aloe plan- 
tations, is in the valleys of these central basins. 

The calcareous strata are markedly unconformable on the 
older rocks. The earlier limestone layers are very hard and 
are darker in color; the more recent ones (fig. iii-7) are simply 
exposed coral reefs and are quite soft and chalky in texture. 
Where exposed, the older limestone (fig. vi-14) erodes into 
characteristic jagged points, separated by irregularly rounded 
holes, which may be several feet in depth; thin slabs of this 
material ring like steel when struck. The central dome has 
been largely removed, but in the middle of Curacao and on 
the southern ends of Aruba and Bonaire, the caleareous rocks 
still practically bridge each island, while capped monadnocks, 
such as the Tafelberg of Sint Hyronimus (Pl. I, Curagao, 
20-N) and Ronde Klip (12-P) on Curacao, show its former 
extent and altitude. Usually the southwestern rim is more 
markedly tilted than the northeastern; the latter is often 
eroded almost to the shore (northern Curacao and Aruba), or 
may remain as flat-topped ridges and mesas of the older lime- 
stone (for example, the coast of Hato, Curacao; fig. v—13), 
while the former is carved by the canyon-like valley outlets 
into bold, angular hills (fig. iii-5).2° Almost invariably, the 
northeastern escarpments of all of the older remnants form 
vertical, or even overhanging cliffs, which are commonly exca- 
vated into more or less extensive caves, and decorated with 


10 K. Martin, I, Plate IX, shows the inland side of the limestone rim 
from Seroe Domi to Seroe Salinja Abau, with the Tafelberg of Santa 
Barbara in the background. 


10 University of Michigan 


stalactites, stalagmites and other seepage deposits (figs. iv—8, 
v-13). In addition, less prominent cliffs may occur above the 
fossil beaches on the leeward side of the hills, and the more 
recent coral layers quite commonly form lower ridges, which 
are separated from the more prominent monadnocks of older 
limestone by valleys parallel to the shore. The true mesas, 
like the Ronde Klip and the Tafelberg of Santa Barbara, are 
almost entirely surrounded by steep cliffs. 

The flora of the islands is discussed in detail by Dr. I. 
Boldingh.? ‘‘The general impression of the vegetation of the 
islarfds Curacao, Aruba and Bonaire is that of a dry country, 
where thorny shrubs and cactuses predominate . . . the vege- 
tation has everywhere a rather uniform aspect. . . . Except- 
ing a few less exposed parts and the higher tops of Curacao 
and Bonaire, the whole vegetation may be said to have a more 
or less xerophile character; in many places where the soil is 
covered by hardly any humus, as on the numerous limestone 
table-lands, it becomes a poor vegetable cover... ; nearly 
everywhere the soil is clearly visible and not covered by a con- 
nected vegetation. . . . The type of vegetation might be gen- 
erally described as a Croton vegetation, . . . determined by 
plants like Croton, Acacia, Lantana, Melochia, Opuntia, Melo- 
cactus. .. . Capparis Breynia is characteristic for the vege- 
tation outside the lime and Rhacoma crossopetalum and 
Antirrhoea acutata for the lime.’’ (Excerpts from pages 
149 and 150.) 

Commonly the omnipresent thorn trees are dwarfed and dis- 
torted by the dry trade winds; the foliage of the divi-divi 
(Caesalpinia coriaria) usually consists of a flat, matted fan, 
which only spreads out to the leeward of the trunk (fig. v-12). 
The giant organ-pipe cacti (Cereus) are among the most con- 
spicuous features of the landscape; planted closely in rows, 
they form many of the fences (fig. vi-14), although the dead 
branches of Acacia tortuosa are as commonly used on Curacao. 
The flat-jointed cacti (Opuntia) are almost everywhere, and 


111914; The Flora of Curacao, Aruba and Bonaire; Leiden. 


Occasional Papers of the Museum of Zoology 11 
may form quite impassable thickets, while the large ‘‘nigger- 
heads’’ of Melocactus and the spiked rosettes of Agave may 
be about the only vegetation near the summits of the larger 
monadnocks of older limestone. 

Nowhere can the vegetation truly be called a forest, 
although, in protected places, especially on the leeward slopes 
of the higher hills, larger trees, such as various species of 
Bursera, Casearia bonairensis, Bumelia obovata, Capparis 
jamaicensis, Machaonia Ottonis, and Guaiacum officinale, do 
form small, or even quite extensive, open groves. However, 
the largest masses of natural foliage are furnished by the 
poisonous ‘‘manzalienja’’ (Hippomane mancinella), which 
forms quite dense and high tree-growths along the dry water- 
courses of the inland region, especially on Curacao. Quite a 
’ few of the shrubs and trees, such as Bursera semiruba, shed 
their leaves during the dry periods, while others, like Guaia- 
cum officinale, are evergreen. 

The native species of molluscs are quite largely confined to 
the limestone outcrops, although certain of them extensively 
penetrate the more heavily wooded valleys of the higher hills, 
especially of those in the Cretaceous rocks (fig. vii-19). At 
the inner borders of the limestone rim (fig. vii-18), the soil 
of the inland region is impregnated with the reddish deposit 
that appears to be the principal residuum from the decomposi- 
tion of coral rocks. Apparently enough calcareous material 
remains in some of these places to support the limestone-loving 
species.1* However, the extensive grass and brush covered 
plains of the central region (fig. vii-17) are practically with- 
out mollusean life. 

The limestone rim is broken by numerous ‘‘roois’’ and, more 
rarely, by deep salt water channels that connect with the 
broad, shallow lagoons behind the shore ramparts. Especially 
on the island of Curacao, the molluscs of each of these lime- 
stone remnants form a quite isolated colony. The result of 
this separation is most noticeable among the forms of the 


12 Included in the ‘‘caleareous soil’’ of Boldingh’s maps. For the 
vegetation of the non-calcareous soils, see page 158, 1. c. 


12 University of Michigan 


genus Tudora on the island of Curacao, where the prominent 
breaks in the limestone rim are accompanied by changes in 
the shells themselves. On Curacao, the main station numbers 
indicate those areas of the limestone which are most markedly 
isolated by such natural boundaries, but, on Aruba and 
Bonaire, sharp lines of demarcation can rarely be drawn.1% 


Curacao 


Although Curacao (Plate I, fig. 1) is only about 35 miles 
long, it can readily be divided into three, quite distinct areas, 
on the basis of the distribution of land molluses. The first 
of these centers about the Tafelberg of Santa Barbara (fig. 
1, 2-B) ; the second includes the remainder of the southwestern 
shore between Spaansche Baai (2—E) and Boca Sint Marie 
(9-G), and the coast of Hato (10, 11, 12) on the northeastern 
side; while the third is composed of the western shore north 
of Bullen Baai (9-F) and the higher hills in the older rocks 
(20). As already discussed, the many breaks in the limestone 
rim of Curacao quite markedly isolate the various stations. 
The more recent limestone is mainly limited to the shore 
region southeast of Piscadera Baai (5-W). 

Station C1. Northwest of New Port (fig. I, 1—-A; L68° 50.72’, 
12° 3.52’).1* New Port is the harbor of the phosphate com- 
pany ; it is situated on a rather broad shelf of recent limestone 
along Fuik Baai. As this locality is on the leeward side of the 


13 Throughout this paper, the stations are designated by letters and 
figures, as follows: C, A, B, and K stand for the islands of Curacao, 
Aruba, Bonaire and Klein-Bonaire, respectively; these, followed by 
b, e, or d indicate (b) the localities on the older deposits, and (c) the 
freshwater and (d) marine habitats, as opposed to the stations on the 
limestone. The figures give the station numbers; these are followed by 
letters for the substations. 

14 After each station is given a reference to the map of the island con- 
cerned (plates I and II), followed by the west longitude and north lati- 
tude of the place mainly collected (indicated by the letter L). These 
are measured to 1/100 of a minute on the Netherlands Government 
1/20000 topographic maps, and thus locate the type localities within ap- 
proximately 20 meters. 


Occasional Papers of the Museum of Zoology 13 


Tafelberg, it is more heavily wooded than are most of the 
stations near the shore. The adjacent slope of the Tafelberg 
is not precipitous and probably similar conditions occur for 
some distance up this side. Near the top, on one of the fossil 
beach levels, occur the rich phosphate deposits that produce 
most of the export from Curagao. 

Station C2. Base of escarpments of the Tafelberg of Santa 
Barbara (2-B; L68° 50.78’, 12° 4.31’ and 68° 51.07’, 12° 4.35’ 
‘for substations C2a and C2b, respectively). This Tafelberg is 
the highest hill (193.8 meters) on the southern portion of 
Curacao, and undoubtedly was a separate island at several 
of the ancient beach levels. The escarpments of the older 
limestone, below the summit of its northern and western sides, 
form almost vertical cliffs which are 200 feet high in places. 
As erosion appears to take place most rapidly at the uncom- 
formity over the underlying, older rocks, these cliffs often 
overhang, and the limestone tends to break off in enormous, 
angular blocks, which slump down the steep slopes below the 
escarpments (compare fig. iv-8). Most of the shells occur in 
the talus from these cliffs; the Tudorae do not extend any 
great distance down the slopes, which are covered with a thin 
soil from the older rocks, but Cerion occurs to the base and 
some distance beyond. These slopes are held by thickets of 
brush, but the larger trees are usually near the base of the 
cliffs. Two localities were collected: the central portion of 
the northern side (C2a) and the base of the western escarp- 
ment (C2b). 

Station Cb2. Manzalienja Rooi, Campo Santa Barbara (2-C; 
L68° 50.63’, 12° 4.94’). A few shells were collected near one 
of the dry water-courses (roois) of the diabasic central region. 
These are commonly bordered by luxuriant groves of large 
trees (Hippomane mancinella), but are usually devoid of 
molluscan life. 

Station C3. Hill north of Fort Beekenberg (3-F; L68° 
52.54’, 12° 4.55’).25 This rather barren block of limestone 


15 For vegetation of nearby crest, see Boldingh, p. 155, ‘‘ Kabrieten- 
berg near Beekenberg.’’ 


14 University of Michigan 


(25.1 meters) just north of the old fort near the Quarantine 
Station, is the type locality of Tudora megacheilos kabrietensis. 
Station C4. Seroe Mansinga (4-H; L68° 53.22’, 12° 4.81’). 
The older limestone remnants between Caracas Baai (3-G) 
and Lagoen Jan Tiel (4-I) reach an altitude of 49.2 meters. 
The brush on the hills themselves is rather dwarfed, but larger 
trees are quite numerous on the northern slopes, just inside of 
the limestone ramparts. In these places, Drymaeus virgulatus 
is especially numerous and occurs even on the lower crotons. 
Station C5. The limestone rim between Lagoen Jan Tiel and 
Sint Anna Baai (5—P) is included in this station. The fol- 
lowing substations represent the main localities collected. 

C5a. Seroe Spanjo (5-K; L68° 54.21’, 12° 5.11’). Very 
similar to the preceding, but most of the collection comes 
from the more recent limestone. 

C5b. Seroe Salinja Abau (5-L; L68° 55.38’, 12° 6.31’). 
Practically a continuation of the Schaarlo. 

C5c. Schaarlo (5—M; L68° 56.55’ 12° 6.81’). Back of Wil- 
lemstad (5—O) is a low ridge (41 meters) of recent limestone. 
It is rather barren, although quite densely covered with 
Croton flavens, ete. Along the northern side, acacias and 
other small thorn trees are more abundant (fig. 11-7). As 
Willemstad is the principal port of the islands, it is probable 
that most of the species described from Curacao by the earlier 
writers came from its near vicinity; and I am regarding the 
Schaarlo as the type locality of all of them that occur there. 

Cid. Fort Nassau (5—N; L68° 56.41’, 12° 7.03’). The old 
fort crowns the summit (68.3 meters) of a triangular monad- 
nock of the older limestone, which is separated from the 
Schaarlo by a small valley. Most of the shells collected come 
from the base of the eastern end of the escarpment, where the 
vegetation is slightly more luxuriant than on the Schaarlo. 

Station C6. This includes the limestone ramparts from Sint 
Anna Baai to Piscadera Baai (5-W). The two main peaks 
of the older limestone, Seroe Pretoe and Veerisberg, together 
with Jack Evertszberg, are known as the ‘‘Three Brothers’’ 
(fig. 1ii-5). Three substations are recognized. 


Occasional Papers of the Museum of Zoology 15 


C6éa. Seroe Quinta (6-S; L68° 56.96’, 12° 6.97’). Behind 
Overzijde, which is the portion of Willemstad on the north- 
west side of the harbor channel, is a ridge (37.7 meters) very 
similar to the Schaarlo. 

C6b. Seroe Domi (6—-T ; L68° 57.57’, 12° 7.32’). This monad- 
nock (84.8 meters) of older limestone bears about the same 
relation to Seroe Quinta that Fort Nassau does to the Schaarlo. 

C6c. Seroe Pretoe (6-U; L68° 58.33’, 12° 7.41’). Black 
Mountain, the first of the Three Brothers, is slightly higher 
(135.5 meters) than the second, Veerisberg (131.5 meters; 
6-V). Most of the collection comes from the leeward slope 
of the former and from the ridge of recent limestone between 
both of them and the ocean. This locality, probably on ac- 
count of the greater altitude of the two peaks, is considerably 
better wooded than the ridges around Willemstad. It is 
separated from Seroe Quinta by a valley in the diabase. 

Station C7. This includes the limestone from Piscadera 
Baai to the Salinja van Sint Michiel (7-B). Two localities 
were collected. 

C7a. Jack Evertszberg (7-Z; L68° 59.39’, 12° 7.92’). The 
lowest of the Three Brothers (115.3 meters) is quite heavily 
wooded along Piscadera Baai and at the base of the northern 
escarpment. Most of the shells obtained come from the ridge 
of recent limestone near the ocean; along this portion the 
sea-cliffs are quite high. I ascended this peak on the day after 
a heavy rain, which had cleared the atmosphere of its usual 
haze (fig. iii-5); from the summit, Bonaire seemed but a 
short distance away, while Aruba could be made out, and the 
higher coastal peaks of Venezuela, from Santa Ana on the 
Paraguana Peninsula to the Cumbres back of Puerto Cabello, 
were plainly visible. 

C7b. Sint Michielsberg (Seroe Blandan; 7—-A; L69° 0.09’, 
12° 9.24’). This peak of the older limestone is quite barren, 
except at the base of the western escarpment, where the col- 
lections were made. 

Station C8. Seroe Spreit (8-D; L69° 0.34’, 12° 9.34’). 
Although this peak of the older limestone, to the north of 


16 Umversity of Michigan 


Salinja van Sint Michiel, is considerably lower (51.2 meters) 
than Seroe Blandan across the lagoon, the western escarp- 
ment is more heavily wooded and has a richer molluscan 
fauna. 

Station C9. Seroe Popchie (9-E; L69° 3.92’, 12° 12.44’). 
This hill of the older limestone (85.3 meters) on the eastern 
side of Boca Sint Marie (9-G) is quite barren, as regards 
both vegetation and mollusecan population. The collection 
comes from the steep, western slope. 

Station C10. Seroe Boca (10-H; L68° 49.19’, 12° 7.14’). 
This low ridge (40 meters) on the eastern side of Sint Joris 
Baai (10-I) is one of the most barren localities collected. The 
cap of older limestone is quite thin and the reddish residuum 
correspondingly prominent. The top of the mesa is quite 
devoid of vegetation, with the exception of two or three giant 
cacti (Cereus). The trade winds, although they come fresh 
from the ocean, are so dry that after a few hours sojourn 
(much of the time in the shade), my conjunctivae became 
so inflamed that I was hardly able to see for some time after- 
ward. However, the deeper crevices of the rocks are green 
with grass, and dwarfed manzalienja trees, which require 
considerable moisture, are bunched on the leeward side of the 
escarpments and detached rocks. The effect of the trades is 
evident, for their tops are leveled off at the limit of protec- 
tion, as if they had been trimmed artificially. The entire 
aspect of the locality gives the impression of a wind-swept 
mountain peak at and above timberline (compare fig. v—11). 
As already mentioned, I suspect that a portion of the moisture 
comes from the heavy dews; I am sure that these form in. 
similar places on Aruba. Despite the vicissitudes of such a 
habitat, Tudora and Cerion are present in the cracks of the 
rocks and even upon the cacti. Peculiarly enough, the speci- 
mens of the former dre larger than the average, although 
those of the latter are much dwarfed (C. wva diablensis). The 
specimens of Cerion from the base of the leeward escarp- 
ment, where they occur even under the manzalienjas, are 
larger than those from the summit of the ridge; the entire lot 
from this station gives a bimodal curve. 


Occasional Papers of the Museum of Zoology 47 


Station C11. This includes four localities along the ridge 
that skirts the coast of Hato. The cap of ancient limestone 
becomes progressively thinner and the vegetation more sparse 
towards the southeastern end, where the conditions approxi- 
mate those of Seroe Boca (C10). 

C11a. Seroe Markita (11-L; L68° 50.95’, 12° 9.06’). This 
lot is from the more barren end (50.1 meters) of the Hato 
ridge. 

C11b. North of Ronde Klip (11—M; L68° 52.06’, 12° 9.38’). 
The southern slope of this portion of the ridge (59 meters) is 
more heavily wooded than Clla. 

C11c. Seroe Papaja (11—-N; L68° 57.20’, 12° 10.71’). The 
dwarfed vegetation on the top of this ridge (62.6 meters) is 
abundant, while the base of the low escarpment, and the 
areas between the large, detached blocks of limestone, are 
grown up with brush and trees and support a thriving mol- 
lusean population. Most of the collection comes from these 
more protected places. 

C1id. Near Landhuis Hato (11-O; L68° 57.92’, 12° 10.95’). 
The high northern escarpment of Seroe Spelonk (70.4 meters) 
is excavated into spacious caves, with enormous stalactites 
pendant from their roofs (fig. v-13). At Landhuis Hato, 
one of the only permanent streams on the islands issues from 
a cave. The nearly level terrane near the base of these cliffs 
supports a luxuriant growth of brush, and the densest stands 
of Cereus that I have ever seen. 

Station C12. Ronde Klip (12—P).*° This mesa is separated 
from the nearby Hato ridge by a valley, in which the diabase 
comes to the surface. It is almost encircled by a high escarp- 
ment of the older limestone, which surmounts the steep, red- 
dish slopes with diabase outcrops. 

C12a. Base of western escarpment. (L68° 52.08’, 12° 9.00’). 
At the base of the limestone cliffs are thickets of brush and 
clumps of larger trees. This is one of the richest habitats 
in central Curacao. 


16 See Boldingh, p. 156; ‘‘ Ronde Klip.’’ 


18 Unversity of Michigan 


C12b. Top of Ronde Klip (LG68° 52.02’, 12° 8.98’). The 
plane summit (129 meters) of the mesa slopes slightly towards 
the north. The brush is dwarfed and much less luxuriant 
than in C12a. 

Station Cb6a. Shore, Campo Marchena (6—X; L68° 57.73’, 
12° 7.72’). An old fruit plantation on the east shore of the 
Schottegat (5-R) was searched, but Succinea gyrata was the 
only species obtained. 

Station Cb6b. Hill with ruins, Campo Blenheim (6-Y ; 
L68° 57.1’, 12° 7.94’). This small knob (20 meters) is formed 
by an outcrop of diabase, but the reddish color of the residual 
soil gives evidence of the former limestone cap (compare 
fig. vii-18). The type locality of Cerion uva desculptum is 
probably some similar hill in this vicinity, as it forms here a 
small proportion of the Cerion population. The shells occur 
only on the rather barren hill, although it is near fertile and 
wooded valleys. 

Station Cb7. Near Landhuis Klein Piscadera (7T-C; L68° 
59.03’, 12° 8.64’). A few dead shells were collected among 
leaves at the edge of an irrigated fruit plantation near the 
highway. Diabase is the underlying rock. 

Station Cb10. Near Seroe Mainsjie (10-K; L68° 49.21’, 
12° 6.75’). This hill (80.7 meters) formed by an outerop of 
diabase, is partially protected by the ridges nearer the shore. 
Cerion uva desculptum occurs here, along with the sculptured 
form. 

Station C18. Seroe Largoe (13-R; L69° 4.18’, 12° 12.17’). 
The sloping top (73.2 meters) of this headland is covered with 
a rather thick growth of brush. At the base of the high north- 
ern and western escarpments of the older limestone, Bursera 
and other large trees are quite numerous. The molluscan 
population of this station is a mixture of the northern and 
central elements of the Curacao fauna. Tudora fossor djeri- 
mensis occurs especially along the base of the escarpment, 
while 7. megacheilos spreitensis and T. muskusi bullenensis 
share the mesa top itself. The cerions are representative of 
the southern subspecies, C. wva wva. 


Occasional Papers of the Museum of Zoology 19 


Station C14. Seroe Grandi (14-V; L69° 7.00’, 12° 14.60’). 
This quadrangular headland (80.2 meters) of the more ancient 
limestone is much more barren of vegetation than are the sta- 
tions farther north, or even C13. The collection comes from 
the northwestern end, and from along the hills just south of 
Sint Jan Baai (14-W). 

Station 015. Seroe di Boca (15—X ; L69° 8.11’, 12° 16.13’) .?” 
This ridge (70.2 meters), just south of Sint Martha Baai 
(15-Y), is capped with a thin layer of the older limestone, 
and the unconformity with older rocks is very evident at the 
base of the northeastern escarpment, where the collection 
(accidentally mixed with one of the lots from C17a) was made. 
The vegetation in this locality is more or less intermediate 
between that of C14 and that of C16. 

Station 016. Seroe Baha So (16—Z; L69° 9.51’, 12° 17.687). 
This lot of specimens comes from the southern escarpment of 
the hill itself (118.2 meters) and from near the edge of the 
shore cliffs (11.4 meters) as far northwest as Sint Kruis Baai 
(16-A). As regards vegetation, this is perhaps the richest 
limestone habitat studied in northern Curagao. 

Station C17. This includes the rim of the older limestone 
formation, from Sint Kruis Baai to Plaja Abau. Three locali- 
ties were collected. 

Station C17a. Seroe Djerimi (17-C ; L69° 9.77’, 12° 20.14’). 
This quadrangular limestone hill (70 meters) is almost entirely 
surrounded by escarpments, which are more precipitous at the 
eastern end (fig. i1i-6). The collections come from the base of 
this escarpment (lot accidentally mixed with that from C15), 
and from the top. 

C17b. Top of shore cliffs (17-D; L69° 10.02’, 12° 20.46’). 
This lot comes from the top of the high shore cliffs (about 10 
meters) between Plaja Djerimi (17-F) and Knip Baai 
(17-D). It is a considerably more barren substation (fig. 
vi-14) than the nearby seroe. 

Ci7c. Sides of Plaja Abau (17-E; L69° 9.83’, 12° 21.24’). 
Plaja Abau ends in a beach of dazzling white, coral sand. 


17 Compare ‘‘ The lime hills near St. Martha,’’ Boldingh, p. 155. 


20 University of Michigan 


On either side, the overhanging shore cliffs (8.2 meters) 
extend for a short distance inland; the calcareous zone at 
their summits is very narrow. The valley mouth itself is 
choked with mangroves and brush. 

Station C18. Limestone remnant (18-K; L69° 9.88’, 12° 
21.41’). North of Plaja Abau, the older limestone formation 
is mainly limited to the short cliffs. A small isolated remnant 
(25 meters) about 400 meters north of Plaja Abau, was col- 
lected during a shower. The rapidity with which the eyclo- 
stomes, and even the cerions, are activated by the rain, is 
astonishing; copulation seems to be the first motive after re- 
lease from aestivation. Peculiarly enough, the specimens of 
Tudora fossor westpuntensis from this station are very small, 
while those of Cerion wva knipensis are among the largest 
collected. 

Station C19. Westpunt (19-M; L69° 10.52’, 12° 22.94’). 
The top (9.9 meters) of the shore cliffs at Westpunt, which is 
northwest of the hamlet of the same name, is bordered with a 
narrow strip of quite barren, older limestone, dirtied by 
abundant pockets of the characteristic reddish residue. Cacti 
form the major portion of the very sparse vegetation. 

Station C20. Tafelberg of Sint Hyronimus (20-N; L69° 
6.90’, 12° 19.32’).18 This high, isolated mesa (229.9 meters), 
more commonly known as Sint Hyronimusberg, is one of the 
most conspicuous landmarks of Curacao. The ancient lime- 
stone cap is thin, so that the encircling escarpments are low, 
but the reddish slopes of the underlying, diabasie rocks are 
also quite steep. The leeward side is well wooded, as is also 
a portion of the top, but the omnipresence of reddish residual 
dust appears to make this a rather barren habitat for land 
molluses. 

Station Cb16. Near Landhuis Sint Kruis (16—-B ; L69° 8.86’, 
12° 18.49’). Southeast of the head of Sint Kruis Baai, an 
outcrop of Cretaceous rocks is crowded with a dense growth 
of organ-pipe cacti. Among these, a few land shells were 
collected. 


18 See Boldingh, p. 157; ‘‘The limestone table-land on Hieronymus- 
berg.’’ 


Occasional Papers of the Museum of Zoology 7 


Stations Cb17 and 20.*° Sint Christoffelberg (372.44 meters) , 
the highest peak in the Dutch Leeward Islands, is the climax 
of a much dissected mass of sedimentary, Cretaceous rocks, 
which outcrop in a complex series of folded and twisted 
laminae. The leeward slopes of these hills, and especially the 
narrow valleys that open to the north and west, are the most 
heavily wooded places in the islands. Especially conspicuous 
are the large clumps of the yellowish-green Bromelia lasiantha, 
which holds water in its sheathing leaves, and may carpet 
areas several acres in extent. In the richer valleys, and 
especially near the top of the western side of Sint Christoffel- 
berg, epiphytes (Tillandsia utriculata, ete.) and lianas (Ficus, 
ete.) increase the luxuriance of the foliage, until it presents 
an aspect very different from the usual arid lowlands of the 
islands. In the broader valleys (fig. 111-6), the divi-divi and 
acacias attain a larger size than elsewhere, and, with the 
organ-pipe cactus and Opuntia, often form quite impassable 
thickets; the larger trees are also more abundant on the hill- 
sides. Peculiarly enough, the land molluses, with the excep- 
tion of Drymaeus virgulatus, bury themselves deeply in the 
rock talus, although the same or similar species may aestivate 
on the brush and trees in much less humid situations. 

Chi7a. Valley between Seroe Bientoe and Seroe Palomba 
(17-C ; L69° 9.47’, 12° 20.29’). This steep-sided, west-facing 
valley between Windy Mt. (224.8 meters) and Dove Mt. 
(163.1 meters) is almost choked with brush (fig. vii-19), but 

_is more barren than the next. 

Cb17b. Valley between Seroes Palomba and Baha Hoendoe 
(L69° 9.10’, 12° 20.34’). This north-facing canyon was col- 
lected during a rainstorm, when the molluses were moving 
around on the surface; for this reason, the shells appeared 
much more abundant than in the other localities examined. 
However, this was the most heavily wooded valley visited. 
Most of the shells were obtained near its mouth, but Drymaeus 


19 Compare Boldingh, p. 157; ‘‘Christoffelberg’’; and K. Martin, I, 
pl. X, XI and XII. 


22 University of Michigan 


virgulatus was more numerous at its head, near the saddle 
between Seroes Palomba and Baha Hoendoe (211.9 meters). 

Cbi7c. Landhuis Knip (17T-H; L69° 9.33’, 12° 20.77’). The 
country house of plantation Knip is on a spur (40.6 meters) 
of Knipberg. A few shells were picked up around the out- 
houses. 

Cbi7d. Irrigated valley (17-1; L69° 9.59’, 12° 21.11’). The 
hofje that ends in Plaja Abau is mainly planted with sorghum, 
but includes a few clumps of fruit trees and bananas. 

Cb20. Sint Christoffelberg (20-P; L69° 8.24’, 12° 20.30’). 
Near the summit of the western side of this hill, the slope is 
very steep, with many small cliffs, but humus and leaves have 
collected on the ledges where trees and brush can gain a foot- 
hold. 


Klein-Curacao 


This low, comma-shaped island (fig. 2) is less than 2 1/2 
kilometers long. The surface is practically flat, and probably 
nowhere attains an altitude of much over 5 meters; I was 
told that the waves break over it during the heavier storms. 
It is very barren and arid, like the eastern shores of the larger 
islands, and the sparse vegetation is entirely composed of 
halophytes. Except for the phosphate deposits on the western 
shore, the bare limestone rocks are exposed everywhere, and 
form low sea-cliffs along the eastern side and at the ends of 
the island. No land molluses were found. 


Aruba 


The island of Aruba is the smallest and most arid of the 
three main islands. The limestone is mainly restricted to the 
southwestern side and the southeastern end, where the more 
recent layers form quite extensive, rather featureless areas, 
which usually slope up quite gently from the low (1 to 2 
meters) shore cliffs. As the older limestone is not so generally 
exposed, high escarpments are rarely formed, although those 
of Seroe Canashito are conspicuous exceptions, while the east- 
ern escarpment near Boca Grandi (fig. I-3, C), and the sides 


Occasional Papers of the Museum of Zoology 23 


of the canyon-like roois around Savaneta (I), are also pre- 
cipitous. The southwestern side of the island is protected by 
a series of coral reefs, which often form narrow islands, less 
than a kilometer off shore (fig. v-12). 

In the diabasic portion of the central region,?° Seroe 
Jamanota (J), the highest hill (188.37 meters) on the island, 
is considerably lower and much less wooded than are the 
higher hills in Curacao and Bonaire. The northern half of 
Aruba has a core of quartz-diorite, which closely resembles the 
rocks of the Pikes Peak Region in Colorado, and also contains 
veins of gold. Enormous, rounded boulders of this material, 
which are often hollowed out on their leeward sides, and out- 
wash fans of angular gravel are conspicuous features of the 
landscape in this. portion.*? 

Although the mollusean fauna changes slightly at Spaansch 
Lagoen (M), Aruba is more nearly a unit than is the case 
with Curacao. 

Station Al. Culebra (A; L69° 52.99’, 12° 25.26’). Near 
the southern end of Aruba is a rolling plain of the older lhme- 
stone, with many sink-holes, which contain the characteristic 
reddish residuum. Most of the shells were collected around 
the buildings of an abandoned mining camp, near the phos- 
phate deposits. Subfossil Cerion uwva is common in this local- 
ity, but no living specimens could be found. The eastern shore 
near Boca Grandi (C) is built up with sand dunes, partially 
held by clumps of ‘‘dreifi’’ trees (Coccoloba uvifera). Most 
of the region is very barren and resembles C10. 

Station A2. This includes the higher plateaus and mesas of 
the ancient limestone, south of Spaansch Lagoen. Most of the 
specimens collected came from three localities. 

A2Za. Near Butucoe (D; L69° 55.44’, 12° 28.23’).22 This 
lot comes from a small, wooded valley near the southern edge 
of Baranca Kasioenti. The higher portions of this plateau are 
about as heavily wooded as any portion of the limestone areas 
in the islands. 


20 See Boldingh, p. 160; ‘‘ Mirlamar.’’ 
21 See K. Martin, II, p. 47, fig. 15. 
22 See Boldingh, p. 159, ‘‘The caleareous table-land near Fontein.’’ 


24 University of Michigan 


A2b. Spur of Seroe Pretoe (E; L69° 54.35’, 12° 26.74’). 
This plateau (altitude about 50 meters), between Roois 
Hundoe and Spoki, is southeast of the main summit of this 
limestone hill. The larger trees are especially common here; 
most of the shells come from around the bases of Guaiacum 
and Bursera trees. 

A2c. Baranca Alto and Isla (F and G; L69° 57.77’, 12° 
28.50’ and 69° 57.45’, 12° 28.28’, respectively).?2 The Isla is 
a rather high, limestone mesa (72.5 meters), which is almost 
completely surrounded by canyon-like roois. Baranca Alto 
(about 50 meters) is the crest at the inner edge of the coral 
formation, along Rooi Taki (K). Both of these localities have 
patches of brush and larger trees, but are considerably more 
barren than the southern substations. 

Station A3. Roo Frances (L; L69° 58.73’, 12° 29.097). 
This lot was obtained from a ledge at the base of the escarp- 
ment of older limestone, on the western side of this rather 
barren canyon. 

Station A4. Seroe Canashito (N). This quadrangular, 
limestone remnant (70 meters) is the only place, north of Rooi 
Frances, where the older limestone develops high, precipitous 
escarpments (fig. iv—8). 

A4a. Base of northern escarpment (70° 0.05’, 12° 30.50’). 
The inner edges of Seroe Canashito are broken off abruptly as 
high, vertical or overhanging cliffs, which surmount the steep 
slopes of the underlying, older rocks. These cliffs contain 
small caves which are decorated with aboriginal picture- 
writing. Most of the specimens come from the talus at the 
base of the northern escarpment. 

A4b. Top of Seroe Canashito (70° 0.07’, 12° 30.44’). 
The sloping top of this mesa is typically eroded into jagged 
points and honeycombed with irregular holes. Most of the 
vegetation is dwarfed and distorted by the wind. 

Station A5.** The exposures of recent, chalky limestone 


23 Compare Boldingh, p. 159; ‘‘The calcareous table-lands near 
Belashi.’’ 
24 Compare Boldingh, p. 159; ‘‘The country near Manshebo.’’ 


= 


Occasional Papers of the Museum of Zoology 25 


along the southwestern shore of Aruba, north of Spaansch 
Lagoen, form gently sloping plains, which are almost com- 
pletely utilized as aloe plantations. However, a narrow strip 
along the top of the low shore cliffs (1 to 2 meters) is too 
jagged and broken by erevices to permit easy cultivation, and 
remains in quite primitive condition. A few distorted divi- 
divi trees and acacias break the monotony of Opuntia and 
dwarfed Jatropha wrens, which does not hide the barren 
rocks (fig. v-12). 

Subfossil shells of the recent species are quite common in 
the aloe fields, along with Oxystyla maracaibensis imitator, 
which was not found alive. Living shells are mainly limited 
to these uncultivated spots, although a few occur within the 
limits of the aloe plantings and even on the pungent, yucca- 
like leaves. 

Ada. Near Perkietenboseh (O; L70° 0.54’ 12° 29.28’). Just 
north of Perkietenboseh, this shore strip is scattered with 
willow brush and small thorn trees. 

Adb. Southeast of Oranjestad (P; L70° 2.04’, 12° 30.52’). 
The more barren portions along the top of the shore cliffs 
necessitate a great deal of work for the net results in shells 
collected. Only a few of the large, flat rocks have any shell 
population on their lower sides, and this mainly consists of 
pupillids. 

Station A6. Limestone remnant, near Tanki Schipau (S; 
L70° 2.0’, 12° 31.88’). A few shells were collected around this 
little mass of the older limestone rocks, in the aloe fields near 
the inner edge of the limestone area. 

Station A7.*> This includes three localities near the north- 
ern end of the island, which is even more arid and barren than 
Seroe Boca (C10) on Curagao. 

A7a. Shore near Malmok (V; L70° 3.33’, 12° 35.93’). In 
this place, narrow strips of recent limestone alternate with 
low, brush-covered dunes. 

A7b. Seroe Annaboet (W; L70° 2.64’, 12° 36.15’). The 


25 Compare Boldingh, p. 160; ‘‘The Hills in the N. W. part.’’ 


26 University of Michigan 


top of this small mesa of the older limestone (25 meters) is 
absolutely barren, but an impenetrable thicket of Opuntia and 
a few, dwarfed thorn trees grow in the shelter of the leeward 
end. A few living shells were collected under the rocks at the 
western end of the top. 

A7c. Seroe Hudishibana (X; L70°, 3.45’ 12° 36.70’). This 
cap of extremely barren, ancient limestone (25 meters) 
is near the northern tip of Aruba. A few dead shells of 
Tudora fossor canashitensis were obtained at the base of the 
leeward escarpment near the lighthouse. 

Station A8. Near Boedoei (Z; L69° 59.35’, 12° 32.89’).¢ 
Along the northern portion of the arid, northeast shore of 
Aruba, the limestone is mainly limited to the immediate vicin- 
ity of the shore cliffs. In a few places, narrow tongues of 
dirty limestone lie in the valleys. Near the gold mines of 
Boedoei, a small limestone remnant (26.2 meters) forms a 
little knob on the northern side of the valley. Although this 
locality is very barren, a few specimens were obtained. 


Bonaire 


Fundamentally, this island is very similar in structure to 
Curacao and Aruba. However, apparently in quite recent 
times, low, almost flat wings have been added on either side of 
the main anticline, the long axis of which runs in the general 
direction of the line between Seroe Brandaris (Plate II-W) 
and the southern Seroe Grandi (I). The northeastern addi- 
tion, which practically coincides with Campo Bolivia (O) is 
the higher (mostly less than 30 meters) ; while the southwest- 
ern one forms the low area (less than 10 meters’ altitude) 
south of Kralendijk. Klein-Bonaire appears to be a continu- 
ation of this latter emergence. Also, the southwestern side of 
the main anticline still retains a large proportion of its cap 
of older limestone, so the entire island has a proportionately 
much greater exposure of the coral formations than either 


26 Boldingh, p. 160; ‘‘The Hills near Andikurie,’’ describes the vege- 
tation of the adjacent outcrops of quartz-diorite. 


Occasional Papers of the Museum of Zoology 27 


Curacao or Aruba. On the whole, Bonaire is better wooded 
than are the other two main islands, but the northern hills do 
not attain the altitude or the richness of those on Curacao. 
uike Aruba, this island has a more homogeneous molluscan 
fauna than is the case with Curacao. 

Station B1. South of Kralendijk (B; L68° 16.68’, 12° 
8.08’).27. The southern shore of Bonaire exposes extensive 
stretches of bare, recent limestone rock (fig. vi-15). Numer- 
ous sink-holes show the presence of fresh and brackish water 
a short distance below the surface; as a result, the vegetation 
is fresh and verdant, wherever it is sufficiently protected from 
the dry winds (fig. vi-16). Most of the shells come from along 
the highway near the western shore. 

Station B2. Shore near Hato (F; L68° 17.51’, 12° 11.49’). 
As Kralendijk is the principal port of the island, the material 
from which the earlier species were described probably came 
from the near vicinity. Everything points to the fact that 
Hartert only collected near the town itself, as he obtained 
Potamopyrgus, which occurs in a well in the town, and the 
small species that live in the more arid situations, but missed 
those from the hills. Neosubulina harterti is the only one of 
his species that I did not find near town, and it doubtlessly 
occurs around some of the buildings. However, it seems pe- 
culiar that he should have missed Cerion uva, which is abun- 
dant everywhere. Hither this station or the preceding one is 
quite representative of the conditions around Kralendijk. The 
present station, along the top of the low shore cliffs north of 
the town, is somewhat richer than B1. 

Station B3. Eastern portion of Montagne. Seroe Largoe 
or Montagne (L) is the main portion of the limestone which 
covers the western side and summit (133.4 meters) of the 
southern portion of the principal anticline. 

B3a. Southeastern escarpment (G; L68° 16.31’, 12° 11.76’). 
The southeastern edge of Seroe Largoe, south of Kibra di Mon- 
tagne, forms a rather prominent escarpment of the older lime- 


27 Compare Boldingh, p. 161; ‘‘The country around Kralendijk.’’ 


28 University of Michigan 


stone. The locality collected is at the base of these cliffs, where 
the vegetation is as rich as in any portion of the windward 
side (fig. iv—10). 

B3b. Kibra di Montagne (H; L68° 16.29’, 12° 12.01’). A 
high saddle (about 100 meters) separates the main hill from 
a narrow limestone ridge that extends in an easterly direction. 
The pass itself is rather barren, although the vegetation is 
much richer in the valleys at either side (fig. iv—9). 

Station B4. Western slope of Montagne (K; L68° 17.24’, 
12° 11.86’).28 The valleys on the leeward slopes of Seroe 
Largoe (Campo Santa Barbara) contain dense thickets of 
brush, interspersed with clumps of larger trees. The station 
is in one of the richer of these, and is perhaps the most heavily 
wooded locality collected on the limestone of any of the islands. 
The crotons attain a height of 2 meters, and form almost im- 
passable thickets, except where the larger trees outshade them. 
Most of the shells come from around the Guaiacum and 
Bursera trees. 

Station B5. Porta Spaio (M; L68° 16.68’, 12° 14.06’).?° 
At the top of the low cliffs that form the northern border of 
the broad plain of Campo Bolivia (O), the larger trees form 
rather open groves, which are as extensive as any in the 
islands. 

Station B6. Base of cliffs near Fontein (P; L68° 17.73’ 
12° 14.66’). The northern escarpment of Seroe Grita Kabai 
(87 meters) is high and precipitous. At Fontein, a tiny rivu- 
let, the only permanent stream on Bonaire, emerges from a 
cave in the older limestone. The locality collected is at the 
base of the cliffs just north of Fontein; the reddish residual 
deposit is especially prominent at this place. 

Station B7. Punta Blanco (R; L68° 12.67’, 12° 10.25’). 
Near the southeastern end of the main anticline, the recent 
limestone, along the eastern shore of the island, is limited to a 
comparatively narrow border. At Punta Blanco (21.5 
meters), the inland edge of this formation presents low escarp- 


28 Compare Boldingh, p. 162; ‘‘The lime table-land near Montagna.’’ 
29 See Boldingh, p. 162; ‘‘On the calcareous soil near Bolivia.’’ 


Occasional Papers of the Museum of Zoology 29 


ments. Near the shore, the surface is very barren (compare 
C10), but at the summit of the leeward cliffs, where this col- 
lection was made, scattered trees indicate a richer environment. 

Station B8. Seroe Wassau (T; L68° 22.58’, 12° 14.187). 
West of the Salinja Goto (U), the older limestone forms a 
rather high hill. The top (123.5 meters) and eastern slopes 
are considerably more barren than B3, 4, 5 and 6, but the base 
of the northern escarpment has developed quite rich vegeta- 
tion. Collections were made in both portions. 

Station B9. Seroe Grandi (V; L68° 21.33’, 12° 17.78’). At 
the northern end of Bonaire, the narrow shore zone of the 
older limestone is excessively barren and arid. In a few 
places, the inner edge rises into hills with steep, southern 
escarpments. The largest of these, Seroe Grandi, protects a 
sparse growth of acacias and organ-pipe cacti. The molluscs 
mainly occur at the base of the escarpment, around and on 
the cacti. 

Station Bb3. Seroe Grandi (I; L68° 16.01’, 12° 11.07’). 
‘*Big Mountain’’ is a rather common name in the Dutch Lee- 
ward Islands. This one is a monadnock (115.3, meters) of 
Cretaceous rocks, near the southeastern end of the main axis 
of the island. The leeward slopes are quite well wooded. This 
is the type locality of Anolis bonairensis Ruthven.*° 

Station Bb5. Traai Montagne (N; L68° 16.35’, 12° 12.78’). 
Near the eastern slope of Seroe Largoe, the somewhat sparser 
vegetation in the region of Cretaceous rocks quite closely re- 
sembles the open groves of the limestone area near Porta 
Spafio (Bd). 

Station Bb7. Inland from Punta Blanco (S; L68° 13.17’, 
12° 10.18’). Behind the protection of the eastern rim, the 
richer valleys of the central region develop thickets of brush 
interspersed with larger trees. Many localities show, by their 
reddish soil, that the erosion of the limestone cap has been 
comparatively recent. 

Station Bb9. Base of Seroe Brandaris (W; L68° 23.97’, 


801923; Occ. Papers Mus. Zool. Univ. Mich., no. 143, pp. 4-5. 


30 University of Michigan 


12° 17.13’).*+ Although extensive thickets of brush and 
clumps of larger trees are present, the northern hills of 
Bonaire are more arid and barren than those of Curacao. The 
few shells collected come from the base of the leeward slopes 
of Seroe Brandaris, the highest peak (240.4 meters) on 
Bonaire. Here the thin soil overlies a substratum of mica- 
porphyrite. 


Klein-Bonare 


Station K1. Klein-Bonaire (Y; L68° 18.02’, 12° 9.51’). 
This island is a low, almost flat mass of recent limestone (6.4 
meters at the highest point), separated from Bonaire by a 
narrow, but rather deep (34 fathoms) channel, which forms 
the harbor of Kralendijk. The general aspect of the surface ~ 
is very similar to that of the region south of Kralendijk (figs. 
vi-15, 16), but the vegetation is somewhat richer, especially 
near the center of the island, where most of the collection was 
made. The smaller shells are most numerous under slabs of 
limestone around the bases of the larger trees. 


FRESHWATER HABITAT 


The permanent bodies of freshwater on the Dutch Leeward 
Islands, fall into three classes: (1) springs and pools in the 
central portion, (2) sink-holes in the recent limestone, and (3) 
the rivulets. As already discussed, the sparse precipitation 
tends to fall in rather heavy storms. During these, and for 
some time afterward, numerous streams pour down the roois, 
and form shallow sheets of water on the flats. In the central 
region of older rocks, extensive systems of artificial dikes hold 
some of this water, and a few of these pools may even outlast 
the dry periods (stations Ce11, 13,17). The porous rocks also 
retain fresh to brackish water just above the level of the salt 
water; this is reached by artificial wells in the older rocks 
(Ce2), and by natural sink-holes in the recent limestone (Bel, 
Kel). All of the permanent streams emerge from caves in the 


31 See Boldingh, p. 163; ‘‘Brandaris.’’ 


Occasional Papers of the Museum of Zoology ol 


older limestone escarpments near the northeastern shores. At 
Fontein van Bonaire, and at Hato van Curacao, these tiny 
rivulets are piped into cement tanks, but at Fontein van Aruba 
(Ac2) a little stream escapes from the roughly dammed 
reservoir at its source. All of this water is heavily charged 
with mineral matter in solution, and a large proportion of the 
stagnant pools are so brackish as to be undrinkable. 

Station Cc2. Reservoir, Campo Wilhelmina (fig. I-1, 2-D; 
L68° 50.57’, 12° 5.38’). In front of Landhuis Wilhelmina a 
small tank is supplied by a windmill. The water is drinkable. 

Station Cc11. Pond, Seroe Papaja (11-N; L68° 57.31’, 
12° 10.63’). In a rooi at the western end of Seroe Papaja, a 
clay dike retains a small, but apparently permanent pool of 
foul, bitter water. Only dead shells were found. 

Station Cc13. Sint Marie Spring (138-S; L69° 4.53’, 
12° 11.86’). Near the western end of Seroe Largoe (13-R), 
and not far from the shore of Salinja Sint Marie (9-G), 
seepage forms a small, but permanent pool of quite fresh 
water. The shells were collected on dead leaves and in the 
algae. 

Station Cc17. Pond, Campo Lagoen (17-L; L69° 9.70’ 
12° 19.83’). The rooi that issues from the valley of Newtown 
has been excavated and walled with stone so that a permanent 
pool is retained. The water is utilized for stock animals. 

Station Ac2. Fontein van Aruba (fig. I-38, H; L69° 54.74’, 
12° 29.54’). On the extremely barren northeastern side of 
Aruba, shoreward from the principal escarpment of the older 
limestone, is an oppressively arid zone of limestone or older 
rock, which, in places, is buried under rather extensive sand- 
dunes. Approach to Fontein from the south reveals what ap- 
pears to be a bank, covered with dead brush, which extends 
from near the escarpment almost to the shore (fig. v-11). Pene- 
tration between this and the cliffs discloses that it is a wind- 
break of living thorn trees, which protects a garden of fruits 
and bananas. The cause of this abrupt change is the Fontein, 
which is roughly dammed at its source under the cliffs, but 
escapes as a small stream, about a foot wide by the same deep, 


32 University of Michigan 


to irrigate the strip of plantation. The shells are very 
numerous on the rocks, in both the reservoir and the brook. 
The water is truly potable. 

Station Bc1. Pos Baca (pl. II, C; L68° 16.32’, 12° 8.44’). 
South of Kralendijk, the low plains of recent limestone con- 
tain numerous sink-holes, which may form small caves; if 
these reach the level of the ground water, they become natural 
wells. The different cavities have a wide range in the saline 
content of the water; in some it is quite fresh, in others dis- 
tinetly brackish, while the salt pans contain highly concen- 
trated sea water. That of Pos Baca (fig. vi-15) is slightly 
brackish, but is used for stock animals, while that of Pos 
Frances, which contains small fishes, is potable. 

Station Kc1. Pos di Cas (Z; L68° 17.56’, 12° 9.70’). Klein- 
Bonaire is very similar in structure to the southern portion of 
Bonaire, and contains several sink-holes which make natural 
wells. One of these forms a small cave near the only habita- 
tion on the island; the water is slightly brackish, but drink- 
able. Analagous cavities are also present on Klein-Curagao ; 
the water in one of these is drunk by goats, but no freshwater 
shells were found. 


NATIVE SPECIES 
NERITIDAE 


Neritina zebra (Bruguiére) 

N. zebra Martens (1879; Chemn., II, 118) ; collected by Deplanche. 

Type locality: Cayenne, Guiana. 

Distribution: Brazil to eastern Venezuela; Panama? Cur- 
acao. I did not obtain this species. Possibly it has been ex- 
terminated by the conversion of many ‘‘salinjas’’ into salt 
pans. 


Theodoxus meleagris (Lamarck) 


Type locality: Rivers of Santo Domingo. 

Distribution: South Carolina, Bermudas and, Mexico, to 
Brazil. Curacao: five dead and one living juvenile specimen 
from the inner end of Sint Kruis Baai (Cd16). 


Occasional Papers of the Museum of Zoology 33 


The radula of the living specimen agrees with this species, 
although it is so juvenile that only 8 points occur on the 
D-lateral. No Neritidae were found in the truly fresh water 
habitats, and these were the only specimens (outside of the 
genus Nerita) obtained from salt or brackish water, although 
a number of promising localities were examined. 


HELICINIDAE 


Stoastomops walkert H. Burrington Baker 
(1924; Naut. XX XVII, 89) 


Type locality: (B4) valley on western slope of Montagne, 
Bonaire. 

Distribution: Bonaire, limestone hills (B3-8) ; Klein-Bonaire, 
central portion (K1). On the underside of surface rocks, 
usually cemented so firmly into cavities and crevices that it is 
very difficult to remove specimens entire. 

Shell (fig. viii-20) : depressed turbinate, subacuminate; thin 
and fragile. Color: golden brown to distinctly reddish. 
Whorls: 414, markedly convex; suture well marked. Later 
whorls: growth wrinkles pronounced but rather irregular; 
spiral thread-riblets fine and numerous (about 30 on last 
whorl), but irregularly spaced, more obscure on base of 
shell. Embryonic shell: 34 whorls; white; practically smooth 
but with irregular growth-wrinkles and punctations present. 
Umbilicus: narrowly rimate. Aperture: subbasal, reniform, 
internally deep orange. Peristome: simple, sharp, incom- 
plete; columellar wall defiected and thickened by whitish 
callus, which spreads out extensively on base of penultimate 
whorl but terminates abruptly, at its distal end, so as to 
form a slight, but distinct emargination just above the basal 
angle. 

Operculum (fig. viii-21): horny plate whitish and very 
thin, but slightly larger than the yellowish, calcareous one; 
the outer surface of the latter very slightly concave, with 
minute, raised punctations; inner surface with eccentric 
growth-lines and with subspiral nucleus at apex of rather 
broad, parietal triangle. 


34 University of Michigan 


Radula (fig. ix-25): B; A/2-3, B/2-3, C/3, D/3, E; 
M 5-6/1, 2-3/2, 1/3, (13-15/3-+ ) 23-25; total 57-61. 
Rhachidian central: broadly shield-shaped with raised, an- 
terior margin. A-—central: somewhat similar to that of 
Stoastoma,** but with two, stout, aculeate cusps and some- 
times a minute accessory one. B-central: similar to A—-central 
but with more ovoid base. C-central: elliptical with 
two, stout, aculeate cusps on the reflected, anterior end, and 
usually a small, outer one. D-lateral: similar to Stoastoma 
but with only two very heavy cusps and a third, vestigial one 
either outside or inside of these. E-lateral: similar to 
Stoastoma but stouter and heavier. Marginals: 23 to 25, of 
which the 5 or 6 inner are unicuspid, 2 or 3 are bicuspid, 1 is 
tricuspid, and the others of the series increase the number of 
cusps rapidly; the outermost uncini are broad and multi- 
cuspid plates as is characteristic in the Helicinidae. 

What I take to be the males are smaller and more de- 
pressed than the dimensions given below. 


Dimensions33 
Shell Aperture Caleareous Opereulum 
altitude majordiam. altitude diameter length width 
2.14 109 (2.34) 42 (.91) 143 (1.31) 44 (.94) 79 (.74) 


This monotypic genus combines, to a remarkable degree, 
the shell and operculum of Eutrochatella, subgenus Pyrgo- 
domus, with the radula of Stoastoma. In general appearance, 
the shell most closely resembles the Cuban species, Z. continua 


321922; Proc. Acad. Nat. Sci. Philadelphia, LXXIV, p. 58, fig. vi-26. 

33 Throughout this paper, the altitude of the shell is given in milli- 
meters, but the other dimensions are expressed as indices, followed by the 
actual measurement (in parentheses). The index of the major or minor 
shell diameters, the altitude of the aperture,or the length of the operculum, 
is the percentage obtained on division by the altitude of the shell. The 
index for the diameter of the aperture is similarly taken in terms of its 
altitude, while that of the operculum is in terms of its length. Unless 
otherwise stated, all dimensions are outside measurements. Altitudes are 
scaled parallel to the central axis of the shell, while the major and minor 
diameters are at right angles to this and to each other. Lengths are 
major dimensions, and widths are measured at right angles to them. 


[Jt] 
on 


Occasional Papers of the Museum of Zoology 


(‘‘Gundlach’’ Poey) and E. pfeifferiana (‘‘Arango’’ Pfr.). 
It differs from all of the species of Pyrgodomus in its narrowly 
rimate umbilicus and the slight but definite notch on the 
columellar wall of the peristome. The incomplete, adnate 
peristome and the Pyrgodomus-like operculum separate Stoas- 
tomops from Stoastoma, with which genus the radula most 
closely allies it. I believe it should be placed in the subfamily 
Helicininae. 


POMATIASIDAE 


Members of this family form the most diversified elements 
of the molluscan fauna of the Dutch Leeward Islands. Most 
of the species are very local in their distribution; the genus 
Cistulops is, as far as is known, entirely confined to these 
islands, while the genus Tudora just reaches the adjacent 
mainland. 

As large numbers of specimens of most of the species and 
subspecies were obtained, an attempt was made to study stat- 
istically their size and shape. For each subspecies, a set of 
shells, if possible from the type locality, was measured for 
altitude and minor diameter. The latter was chosen as less 
variable and more easily measured than the major diameter. 
As the shells were almost universally decollated, it was also 
necessary to count the whorls retained; quarter-whorls were 
taken as unit differences. From these dimensions, the minor 
diameter index was obtained. As the sexes of all of the 
specimens had not been noted during removal of the animal, 
a curve was made by counting the individuals of each minor 
diameter (in tenths of millimeters). In all cases where the 
specimens had come from a single locality, this curve was 
bimodal; and the lowest internodal point was taken as the 
dividing line between the two sexes. The obvious error, intro- 
duced by this approximation, was checked by reference to the 
modes and by comparison with the curves of species of 
Tudorata, in which the sexes do not intergrade in size; I be- 
lieve it to be practically negligible in most cases, but it in- 
creases in the smaller species. Using this line of separation, 


36 University of Michigan 


the shells were arranged in columns, according to sex and 
number of whorls, and the means for altitude, minor diameter 
and index were computed for each class, and for the total 
population of each sex. 

In the tables of dimensions (tables IV—VII) are presented 
the data for the entire population and for some one of the 
whorl classes. As might be expected, the mean altitude in- 
creases and the mean minor diameter index decreases with 
the number of whorls retained. The males retain a smaller 
number of whorls than do the females; this appears to be a 
matter of sex and not of comparative size, as the mean dia- 
meter is usually quite constant in those whorl-classes near the 
mean, although very small shells often retain a smaller number © 
of whorls than do the very large ones of the same sex. In 
Cistulops ravem arubana, complete individuals of both sexes 
were obtained, and the females appeared to develop a greater 
number of whorls than did the males. In the larger species 
studied, the males are slightly more slender than the females 
with the same number of retained whorls; this also may mean 
that the males tend to develop a lesser number of whorls. 
Complete males were obtained of Cistulops ravem raveni, 
Tudora maculata, Tudorw fossor fossor, and Tudora fossor 
canashitensis ; the data for these are given. In the other sub- 
species, the number of whorls is that estimated by the com- 
parison of young shells and modal, decollated females; I sus- 
pect that the male would average 144 to 1 whorl less. The 
rather large variation in all dimensions is probably due, in 
large part, to variation in the number of whorls, which appears 
to be rather characteristic of these desert shells (compare 
Cerion uva). The apparent variation is also partially 
caused by the rather large whorl-classes chosen; data from a 
single whorl-class thus give a markedly flat-topped curve. For 
these reasons, apparent intergradation, in the indices of two 
species, does not necessarily indicate actual intergradation in 
shell form. 


Occasional Papers of the Museum of Zoology 37 


CISTULOPSINAE34 


Cistulops raven raven (Crosse) 


Cistula raveni Crosse (1872; J. de C., XX, 159); Crosse and Bland 
(1873; J. de C., XXI, fig. I-5). Chondropoma (s.s.) raveni Henderson 
and Bartsch (1920; P. U. S. Nat. Museum, LVITI, 62). 


Type locality: ‘‘Curacao’’; probably the Schaarlo (C5c), 
back of Willemstad; collected by Raven. 

Distribution of species: Curacao; north of Seroe Salinja 
Abau and Ronde Klip; Aruba, south of Rooi Frances. 764 
specimens collected. 

Distribution of subspecies: Curacao; Seroe Salinja Abau 
and Ronde Klip, north of Plaja Abau (C5, 6, 11, 12, 15, 17, 
20). Rare to quite common; buried rather deeply under rocks 
and in crevices in talus, on the more humid, wooded portions 
of the limestone mesas and escarpments. <A purely terrestrial 
species. 343 specimens collected. 

Shell: small, slender obovate, thin. Color: cream to brown- 
ish, with 7 or 8, chocolate-colored spiral bands, usually broken 
into rows of flammulations except on the last 14 whorl. 
Whorls: 7 and 81% in two complete males; about 414 
retained; not markedly convex, and with quite shallow sutures, 
at least on the middle whorls; last 1% whorl markedly de- 
scends, decreases in diameter and is solute and angulate above. 
Last whorl: growth sculpture of microscopic, undulating, in- 
complete and anastomosing wrinkles, and high, compressed, 
widely-spaced riblets, which are crested at the upper angle 
or form prominent buttresses at the suture; riblets much more 
closely spaced near peristome; spiral sculpture of numerous 
(15-18 counted), low, rounded cords, which are higher and 
more prominent in the umbilical region, where the growth 
riblets form cusp-like crests over them. Earlier whorls: 
similar, but spiral sculpture less prominent, and microscopic 
growth-wrinkles slightly more regular. Embryonic whorls :*° 
134; practically smooth, but very slightly and irregularly 


34 See 1924; Naut., XXXVII, 89-92. 
35 1923; Occ. Papers Mus. Zool. Univ. Mich., no. 137, fig. I-6. 


38 University of Michigan 


wrinkled; next whorl with microscopic growth-wrinkles and 
numerous, low growth-ridges; spiral sculpture and definite 
growth riblets begin on 3rd whorl; all apical whorls convexly 
rounded and with well-impressed sutures. Umbilicus: small, 
tubular; hidden by last whorl. Aperture: broadly obovate, 
almost circular, and with long axis at about 45° to that of the 
shell; internally light-colored with prominent, spiral bands. 
Peristome: duplex; outer portion abruptly deflected; parietal 
angle with prominent, lanceolate auriculation, which is con- 
cave externally and is slightly twisted backwards at its inner 
edge; lower palatal, basal and lower columellar walls with 
(almost invariably) 5 elliptical reflections, separated by 6 
deep sinuses which correspond to the positions of the more 
prominent color bands; upper columellar region broadly re- 
flected; parietal wall narrowly reflected; inner portion of 
peristome continuous but without internal callus. 

Operculum (fig. xi-42): 4 to 414 whorls; almost circular; 
nucleus subcentral but slightly nearer basal angle; chondroid 
plate heavy, almost flat but with rim of each whorl slightly 
raised ; calcareous portion represented only by irregular punce- 
tations. 

Radula (fig. ix-26): C/3; L1/6-8 + 1/4; M/28-32. Rha- 
chidia central: cowl-shaped, with large cusp curved over the 
anterior end and a minor one on each side; thickened portion 
of base ovoid, but thinner wings make the entire tooth a 
truncated isosceles triangle. Inner lateral: shape similar to 
central but asymmetrical, with the thickened, inner edge 
longer than the thin, outer one; the three principal cusps are 
similar in arrangement to those of the central, but the middle 
one almost always bears two serrations on either side near its 
base; on a few teeth only one of these last could be seen on 
the inner side while in a few others three were counted on the 
outer side (the refraction of the edge of the main cusp makes 
the definition of these small points very poor). Outer lateral: 
with a curved, rectangular base which bears 4 stout, aculeate 
cusps at its outer end. All of the three inner teeth are thick- 
ened transversely just below the cusped tips; when the second 


Occasional Papers of the Museum of Zoology 39 


lateral is viewed in profile (view labeled 2 in figure), this 
shows as a blunt, rounded projection. Marginal: with a 
series of recurved, lanceolate cusps, which decrease in size 
towards the outer edge; the edges of the outer ones pectinate 


Measurements : 
Shell Peristome Aperture Whorls 
alt. maj.diam. min.diam. alt. diam. inside 


Crosse and Bland; 

type, from figure 7.0 47(3.3) 34(2.4) 103(2.4) 22(1.5) 4 
C12, male; complete 8.8 40(3.5) 31(2.7) 30(2.6) 96(2.5) 17(1.5) 8% 
C15, male; complete 7.2 44(3.2) 35(2.5) 32(2.3) 96(2.2) 19(1.4) 7 


C5. Living specimens rare (19 collected). 

C6. Very infrequent (12 specimens). 

Clicd. Frequent (84 specimens). 

C12. Frequent (42 specimens). 

C15. Quite common (82 specimens). One albino female. 
C17. Quite common (94 specimens). 

C20. Quite infrequent (10 specimens). 


Table IV. Dimensions of Cistulops raveni and Tudora maculata 


Subspecies ravem arubana maculata 
Locality C11 A2e B5 
Number of individuals; males 41 98 30 
females 38 40 21 
Mean and extremes of 
whorls retained; males 4.2(4.0-5.0) 4.1(3.8-4.5) 3.8(3.3-4.3) 
females 4.3(3.8-5.0) 4.2(3.8-5.0) 3.9(3.5-4.3) 
Mean altitude; males 6.7 7.2 7.0 
females 7.7 8.4 7.8 
Mean index (minor di- 
ameter) ; males 40 40 43 
females 38 39 42 
Mean and extremes of 
minor diameter ; males 2.7(2.5-2.8) 2.9(2.5-3.1) 3.0(2.9-3.1) 


females 2.9(2.8-3.1) 3.2(3.1-3.5) 3.3(3.2-3.5) 

Mean and extremes of 
altitude (4 whorls); males 6.6(6.1-7.0) 7.2(6.5-7.7) 7.1(6.7-7.6) 
females 7.4(7.1-7.7) 8.0(7.7-8.7) 7.8(7.6-8.0) 

Mean and extremes of 
index (4 whorls); males 40(39-42) 40(38-42) 42 (41-45) 
females 40 (38-41) 39(38-41) 41 (40-43) 


40 University of Michigan 


the distal edge of the tooth to almost 1/3 of its depth; laterad 
to the cusps, fully half of the distal edge of the entire tooth 
is undivided, slightly thickened and recurved. 

Animal: very dark in color, almost black; foot very short, 
almost circular; snout long; tentacles stout, clavate. 


Cistulops raveni arubana, new subspecies 


Type locality: (A2b) spur of Seroe Pretoe, between Roois 
Spoki and Hundoe, Aruba. 

Distribution: Aruba, Rooi Frances, south to Seroe Pretoe 
and east of Fontein (A2, 3). Infrequent to common; habitat 
similar to ravent. 421 specimens collected. 

Shell: males about as large as females of ravent. Color: 
typically albino, but more commonly as in ravent. Whaorls: 
74% in an entire male; 834 and 9 in two entire females. 
Last whorl: growth riblets commonly thinner and more widely 
spaced than in raveni. Other characters as in raveni. 

Operculum: similar to raveni. 

Radula (fig. ix—26, no. 2): very similar to raveni, but the 
minor serrations of the central cusp of the inner lateral appear 
somewhat heavier. 


Measurements 
Shell Peristome Aperture Whorls 
alt. maj.diam. min.diam. alt. diam. inside 


A2,male; complete 8.2 39(3.2) 33(2.7) 32(2.6) 100(2.6) 18(1.5) 7% 
female; entire 10.5 35(3.7) 31(3.2) 28(2.9) 104(3.0) 17(1.8) 8% 
female (type) 11.1 33(3.7) 31(3.4) 26(2.9) 107(3.1) 16(1.8) 9 


A2. Quite common (407 specimens collected). 28 specimens are quite 
unicolor, white to cream. The shells from the southern stations (A2a, b) 
average even larger than the lot from A2e. 

A3c. Infrequent (14 specimens). 


Although all of the lots of typical raveni average noticeably 
smaller than those of arubana, the two forms intergrade ex- 
tensively. The much larger percentage of albinos (the typical 
form) on Aruba also seems evidence of racial differentiation. 


Occasional Papers of the’ Museum of Zoology 41 


CHONDROPOMINAE 


Genus Tudora 


Tudora Gray (1850; Brit. Mus. Cat. Cycloph., 48); monotype Cyclos- 
toma simile ‘‘Gray’’ Sowerby. 

Shell: ovate conic to elongate ovate. Sculpture: exceed- 
ingly variable, but the growth ribs are not tufted at the suture 
nor is the latter channeled. Embryonic whorls: 114, not 
distinctly limited; cream to dark horn-colored, commonly 
darker below; practically smooth, but irregularly and very 
minutely wrinkled. Peristome: simplex, sharp; incomplete or 
continuous; not abruptly reflected. 

Operculum: perimeter channeled; chondroid plate rela- 
tively thin; calcareous portion consists of vertical growth- 
lamellae, which are often cemented together by interstitial 
material, and which coalesce at their distal edges to form a 
caleareous plate; the last is almost parallel to the chondroid 
one, and is marked externally by rather weak growth-wrinkles. 

Radula: central and inner lateral with heavy, triangular 
bases, each with a single, heavy cusp; outer lateral with rec- 
tangular base and markedly reflected tip, which bears about 
five aculeate to spatulate cusps; marginal with numerous, 
recurved, lanceolate cusps, which fill more than half of the 
entire distal edge, outer uncusped portion abruptly sloping 
down to relatively narrow base. 

Although the radulae of the Chondropominae are simple 
and rather uniform in structure throughout the subfamily, 
they do show characters of considerable systematic value, 
which I hope to present in a future paper. Amongst many 
others, I have examined the radulae of several of the species 
that belong to the Jamaican, Mexican and Central American 
group, usually known as Colobostylus, and believe it to be 
quite distinct from Tudora. In addition, the simplex, slightly 
expanded peristome of the latter genus is quite different from 
the duplex, thickened condition in the northern group, and 
was utilized for their separation by most of the earlier writers. 
I have not been able to examine the radula of any of the 
Cuban and Haitian species, which include typical Licina, but 


42 University of Michigan 


which were placed in Tudora s. s. by Henderson and Bartsch 
(1920, p. 77). However, the peristome of these species appears 
to be more like that of Colobostylus, and it seems probable that 
they will be found to be more closely related to some of the 
other groups of the Greater Antilles. In this connection, 
attention is called to the fact that Licina ‘‘Browne’’ Gray 
(1847; P. Z. S., XV, 81), monotype ‘‘Turbo labea,’’ is three 
years prior to Tudora Gray (1850). In the present paper, 
the genus Tudora is considered to include only the species 
listed below under the three minor groups, Bonairea, Tudorata 
and Tudoras.s. Regardless of the question of generic inclusion, 
these three groups are certainly quite distinct from any of the 
other groups of American Pomatiasidae, although Tudorata 
possesses certain characters in common with a mainland group, 
which includes Adamsiella aripensis Guppy, from Trinidad, 
Cyclostoma aspratilis Morelet, from Ecuador, Cyclostoma 
rigidula Morelet, from Guatemala, and my ‘‘ Tudora’’ william- 
som, from Venezuela. 


Subgenus Bonairea, new 


This subgenus is characterized by the small size, the solute 
last whorl, the simple, continuous, unreflected peristome, the 
convex operculum (quite similar to Ramsdenia), and the stout, 
aculeate cusps of the outer lateral tooth of the radula. The 
chondroid plate of the operculum is smaller than the interior 
of the aperture, but the calcareous plate is larger; the entire 
structure fits the peristome like a flanged cover and the animal 
is usually unable to draw it inside. In all of the other species 
of Tudora, as in most of the Chondropominae, the operculum 
is readily withdrawn into the aperture. The radula of 
Bonairea is quite different from that of Ramsdenia, which is 
closely related to other Greater Antillean groups (Annularella, 
Rolleia, and Blaesospira). Monotype: Zudora maculata. 


Tudora maculata ‘‘Bland’’ H. Burrington Baker 
(1924; Naut. XX XVII, 92) 


Type locality: (B8a) base of east-facing escarpment, about 
Y% kilometer south of Kibraé di Montagne, Bonaire. 


Occasional Papers of the Museum of Zoology 43 


Distribution: Bonaire; the higher limestone hills, from 
Montagne and Seroe Largoe north to Fontein and east (at 
least) to Seroe Wassau (B3, 4, 5, 6, 8). In the more humid, 
wooded places; buried rather deeply under rocks and in the 
erevices of the talus. - 175 adults collected. 

Shell (fig. vili-22) : small, elongate-conie, thin. Color: light 
to dark brown, with about 8, chocolate-colored, spiral bands, 
usually broken into rows of flammulations. Whorls: 644 (in 
a complete male), of which a, little less than 4 are usually 
retained; cylindrical, with very deeply impressed sutures; 
last 14 whorl descending, slightly tangential, gradually con- 
stricted, and solute. Last whorl: growth sculpture of crowded, 
regular, low, rounded threads, interspersed with more promi- 
nent riblets, at intervals which appear to correspond to resting 
periods as they are crowded closely together near the per- 
istome; spiral sculpture only visible under lens, not especially 
prominent in umbilical region, consisting of widely-spaced, 
fine threadlets, which sometimes crenulate the growth threads, 
but are more commonly obscured by them. Earlier whorls: 
sculpture somewhat more prominent. Umbilicus: small, tubu- 
lar. Aperture: circular; internally light colored. Peristome: 
simple, continuous; scarcely, if at all, thickened. 

Operculum (figs. vili-23, 24): 414 whorls; almost cirecu- 
lar; nucleus subcentral, but slightly nearer the basal angle; 
chondroid plate inconspicuous, markedly concave internally ; 
calcareous portion thick, solid, and with channeled perimeter; 
external surface (caleareous plate) distinctly convex, larger 
than the chondroid plate, and marked by fairly prominent 
growth-wrinkles. 

Radula (fig. xi40): C/1:L1/1+ 1/5; M/40-41. Central: 
base very heavy, forming a truncated, isosceles triangle ; single 
cusp heavy but slender, almost aculeate. Inner lateral: similar 
to central but asymmetrical, with inner side longer than outer. 
Outer lateral: with long, rectangular base; anterior reflection 
with 5 stout, aculeate cusps; the transverse thickening under 
the reflection is very heavy, so that it produces a marked 
angulation of the sides of the base (as viewed in profile). 
Marginal: relatively heavier than in Tudora s. s. 


44 Unversity of Michigan 


Measurements 


Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male’ 7.4 53(3.9) 42(3.1) 27(2.0) 95(1.9) 27(2.0) 90(1.8) 4 
female 
(type) 88 51(4.5) 41(3.6) 27(2.4) 96(2.3) 26(2.3) 91(2.1) 4 


B3. Infrequent (9 specimens collected). The type locality. 

B4. Quite common (84 adults). Shells larger than those from B5. 
B5. Frequent (51 adults). Lot measured (see Table IV). 

B6. Quite infrequent (15 adults). Large, light colored. 

B8. Quite infrequent (16 adults). Like Bd in size. 


Subgenus Tudora s. s. 


Shell: larger and heavier than in Bonairea. Post-embryonic 
whorl : horn-colored, with fine, regular and quite closely-spaced 
growth-riblets; all apical whorls high, convexly rounded and 
with well-impressed sutures. Aperture: ovoid. Peristome: 
incomplete to almost continuous; sharp; simply thickened or 
expanded. 

Operculum: ovoid; nucleus markedly eccentric; external 
surface of calcareous plate almost flat to concave. 

Radula: cusps of outer lateral are simply spatulate lobes 
of reflected portion. 


Section Tudorata, new 


Shell: elongate-conic. Males markedly smaller than the 
females; the two sexes scarcely intergrade in size. Sculpture: 
very variable, but always with a marked tendency to accen- 
tuate the spiral cords in the umbilical region. Post-embryonie 
whorls: color bands and spiral sculpture begin at about the 
fourth whorl. Peristome: sharp; simple, or gradually ex- 
panded in the palatal and basal regions but abruptly narrowed 
in the columellar. 

Operculum: chondroid plate slightly concave internally; 
ealeareous plate rather thin, much smaller than the horny 
base; supporting lamellae not greatly thickened and usually 
incomplete, so as to leave a considerable suleus around the 
outer edge of the caleareous portion. 


Occasional Papers of the Museum of Zoology 45 


Type species: Zudora muskust. Besides the three species 
on Curacao and Bonaire, this group apparently includes 
Tudora costata (‘‘Menke’’ Pfr.), habitat unknown, and 
Tudora plicatula (Pfr.), from Puerto Cabello, Venezuela.*® 
All of the species are more or less arboreal. 


Tudora aurantia aurantia (Wood) 


Turbo aurantius Wood (1828; Index SuppL, fig. vi-23) ; compare Smith 
(1898; Proc. Mal. Soe., III, 116). Cyclostoma awrantiacum Sowerby 
(1843; Thes., fig. xxiv-46, 47), not Deshayes (1834; Bélanger voy. Zool., 
146). C. versicolor Pfr. (1846; Zeit. Mal.,33; and 1847; Chemn., IT, fig. 
ix-13, 14) ; substituted for preceding on account of Annularia aurantiaca 
Schumacher (1817). C. carnewm ‘‘Menke’’ Pfr. (1847; Chemn., IT, p. 
65, fig. ix-1l, 12); more cancellate form. Tudora versicolor Bland 
(1868; Amer. J. C., IV, 192); the first Bonaire record. 


Type locality: unknown; probably in the vicinity of Kra- 
lendijk, Bonaire (B2). 

Distribution of species: Bonaire and Klein-Bonaire; every- 
where on limestone outcrops, on trees and brush and under 
rocks; also occurs in the more heavily wooded localities, in the 
region of the older rocks. 1,378 specimens collected. 

Distribution of subspecies: Bonaire ; southern portion, north 
to Porta Spafio and Seroe Montagne (B1-5; Bb3, 7). 781 
specimens collected. 

Shell (figs. xii-A): solid. Ground color: cream to pink; 
spiral bands present or absent, orange to chocolate, broken 
or entire; eroded apex and apical plug scarlet and orange to 
deep indigo, almost black. Whorls: about 914, of which 
43/, are usually retained; not markedly convex and with 
suture shallow but distinctly marked; gradually increasing 
in diameter from first to last. Seulpture of last whorl: growth- 
cords quite regular, low, broadly rounded and usually closely 
spaced; spiral sculpture obscure (aurantia) except in umbili- 
cal region, to almost as prominent as growth-sculpture (car- 
nea). Earlier whorls: growth sculpture more delicate, so that 


36 1923; Occ. Papers Mus. Zool. Univ. Mich., no. 137; pp. 23-26, fig. 
IT-9. 


46 University of Michigan 


the spiral sculpture is conspicuous. Third and fourth whorls 
(young shells): with delicate but high, slightly undulate, 
widely-spaced, growth costulation, which forms sutural but- 
tresses distinctly reminiscent of 7’. plicatula; spiral sculpture 
indistinct. Umbilicus: subrimate, more open than in rupis or 
muskusi. Aperture: ovoid, with long axis almost parallel to 
that of the shell; internally light buff to deep orange, bordered 
with white callus. Peristome: simple, sharp, almost or quite 
entire; very slightly produced into parietal angle, where it is 
commonly double; thickened internally by white, rounded 
eallus, which is always continuous in adult shells. 

Operculum: 314 whorls: caleareous plate heavy (for the 
section) and distant from the horny base; outer surface con- 
cave at the nucleus, but with the last whorl quite markedly 
convex along a line parallel to the palatal edge. 

Radula: quite similar to muskusi. 


Measurements 
Shell Aperture Whorls 
alt. maj. diam. alt. diam. 


aurantius Wood 

from legend (1828) 12.7-19.0 

from figure (1828) 14.9 53( 7.9) 37(5.5) 80(44) 5% 
aurantiacum Sowerby 

from figure (1843) 18.1 49( 88) 37(6.6) 77(5.1) 5% 
versicolor Pfr. 

from figure (1846-9) 17.1 57( 9.8) 38(6.5)  83(5.4) 4% 
carneum Pfr. 

from figure (1846-9) 16.7 66(11.0) 438(7.2) 78(5.6) 4%4 


B2. Quite abundant (195 specimens collected). As it is very probable 
that the vicinity of Kralendijk is the type locality of this species, these 
specimens, from near the shore just north of town, are taken as topo- 
typical material (see Table V). 

Bl. Common (134 specimens). Very similar to the preceding lot, but 
with the sculpture of the last whorl even more reduced. In some speci- 
mens, the growth threads are so broad and low as to coalesce and render 
the surface of the last whorl almost smooth. 

3a. Abundant (105 specimens). Very similar to B2, but attain a 
somewhat greater size. 

B4. Abundant (32 specimens). Very similar to B3a. 

B5. Very common (19 specimens). Very similar to B1. 


Occasional Papers of the Museum of Zoology 47 


B7. Common along the western escarpment (203 specimens). Simi- 
lar to Bl, but considerably heavier; the peristomal callus is especially 
well developed. Usually more bleached than the other lots, but one speci- 
men is solid orange and another dark purplish-brown. 

Bb3. Frequent (55 specimens). Darker and more strikingly colored. 
Growth threads finer, and practically all of the shells markedly cancellate. 

Bb7. Quite infrequent, even in the richest portions (38 specimens). 
Similar to B1; some specimens are so smooth as to show a polish. 


Table V. Dimensions of Tudora aurantia, T. rupis and T. muskusi3? 


Wassau- newport- bullen-  grandi- 
Subspecies ensis aurantia — ensis rupis ensis ensis muskusi 
Locality B8 B2 Cl C2 C13 C14 Cl17be 
Number of 

individuals 115 90 105 48 26 146 74 
140 105 95 47 28 164 91 

Mean whorls 

retained 4.5 4.6 4.8 4.6 5.3 4.7 4.8 
4.8 4.8 4.9 4.7 5.7 5.2 5.3 


Extremes, 
ditto 4.0-5.3 43-60 43-63 4.0-63 5.0-6.0 4.3-6.0 4.3-6.0 
43-55 43-55 438-5.5 43-53 43-65 45-65 43-58 


Mean alt. 12.8 12.7 13.2 ES 12.1 11.0 11.9 
17.0 16.4 AST 13.2 GaL 14.9 15.9 

Mean minor 
diam.index 46 48 47 47 44 46 47 
47 49 47 48 45 47 48 

Mean minor 
diameter 6.0 6.1 6.2 5.3 5.3 5.2 5.6 
8.1 7.9 7.4 6.3 7.2 7.0 7.6 

Extremes, 


ditto 5.2-6.9 5468  5.3-6.7 4.6-5.7 4.7-6.1 46-58  5.0-6.3 
7.2-9.1 749.0 6.9-8.0 5.8-7.9 6.7-7.9 647.7 7.0-8.3 


Mean alt., 
5 whorls 13.6 13.1 13.3 11.8 11.6 11.6 12.2 
17.4 w7.0 15.9 13.7 14.7 14.5 15.6 

Extremes, 


ditto 12.2-14.9 12.2-13.5 11.5-14.1 10.2-11.9 10.8-12.1 10.7-12.3 11.3-12.8 
15.6-18.8 15.3-18.2 14.7-17.2 12.8-14.9 14.7 13.4-15.4 14.7-16.2 
Mean index, 


5 whorls 45 47 46 46 45 45 47 
47 47 47 48 47 48 49 
Extremes, 
ditto 42-47 46-49 43-50 45-48 44-48 42-47 45-49 
4449 44-50 45-49 46-50 47 46-51 47-52 


37 Males in upper horizontal row under each heading; females in lower. 


48 University of Michigan 


Tudora aurantia wassauensis, new subspecies 


Type locality: (B8) Seroe Wassau, just west of entrance to 
the Goto, Bonaire. 

Distribution: Bonaire; northern portion, south to the Goto 
(at least) and east to the cliffs near Fontein (B6, 8, 9; Bb9). 
Klein-Bonaire: (K1). Quite abundant in similar habitats to 
aurantia, also in favorable valleys in the higher hills of the 
older rocks (Bb9). 597 specimens collected. 

Shell (fig. xii-B): heavier and slightly larger and more 
slender than aurantia. Growth seulpture (fig. x-28): con- 
siderably heavier, higher, and more widely spaced, so as to 
expose the distinctly marked spiral sculpture. Peristomal 
callus: heavier. Other characters as in aurantia. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 13.5 56( 7.5) 45(6.1) 40(5.4) 78(4.2) 31(4.2) 78(3.38) 5 
female 18.8 56(10.5) 47(8.8) 39(7.4) 80(5.9) 28(5.3) 85(4.5) 514 
female 

(type) 18.2 57(10.4) 46(8.4) 40(7.2) 83(6.0) 29(5.3) 85(4.5) 5% 


. B8. Quite abundant (155 specimens collected). Typical lot (see 
Table V). 

B6. Very common (13 specimens). Somewhat smaller. 

B9. Very infrequent; mainly on Cereus and under rocks at the base 
of these cacti. Bb9. Rare (141 specimens altogether). Somewhat 
smaller. The specimens from Seroe Grandi (B9) are bleached, and quite 
commonly stained with a rusty deposit, as appears characteristic of speci- 
mens of Tudora from these wind-swept, excessively arid places. Growth 
sculpture widely spaced but lower; scarcely more prominent than the 
spiral; the entire shell distinctly cancellate. 

Kl. Very common (288 specimens). Large and heavy; somewhat 
bleached. Altogether more or less intermediate between wassauensis and 
aurantia, this lot from Klein-Bonaire is considerably closer to thé for- 
mer; this is especially peculiar as the conditions are similar to those in 
the type locality of awrantia, just across the harbor. 


Although this subspecies intergrades with typical aurantia, 
large lots give an impression of a markedly more costate shell. 
In the typical lot of wassauensis, the growth sculpture is rela- 


Occasional Papers of the Museum of Zoology 49 


tively much more prominent than the spiral sculpture, 
although the latter is usually more evident than in aurantia. 


Tudora rupis rupis H. Burrington Baker 
(1924; Naut. XXXVII, 93) 


Type locality: (C2a) at base of northern cliffs of Tafelberg, 
of Santa Barbara, southern Curacao. 

Distribution of species: Curacao; limestone of western side, 
south of Spaansche Baai. 290 specimens collected. 

Distribution of subspecies: Curacao; only found near the 
base of the cliffs on the northern and western sides of the 
Tafelberg (C2). Quite common; on trees and brush, also 
found under limestone rocks. 95 specimens collected. 

Shell (fig. xii-E): somewhat less solid than aurantia. 
Ground color: white to buff or pink; spiral bands present or 
absent, pink to very dark chocolate with purplish tinge, rarely 
solid and then usually coalescent, normally broken and com- 
monly with blotches connected so as to form irregular varices 
parallel to the growth-sculpture ; eroded apex and apical plug, 
orange to purplish black. Whorls: about 9, of which about 
414 are usually retained; not markedly convex and with 
suture even shallower than in aurantia. Last whorl (fig. 
x-29): growth-cords quite regular but usually obscure, low 
and rounded; spiral sculpture of about 8, prominent, rounded 
ridges, which (typically) are scarcely surmounted and not 
broken by the growth sculpture. Earlier whorls: growth 
sculpture relatively more prominent; either cancellate or 
with spiral sculpture surmounted by ecusp-like thickenings 
of the growth sculpture. Third and fourth whorls (young 
shells): with more widely spaced and stouter growth-riblets 
than in aurantia; these show less tendency to form buttresses 
at the sutures. Umbilicus: rimate. Aperture: ovoid, with 
long axis quite oblique to that of shell; internally buff to 
chocolate-brown, and sometimes showing the spiral bands; 
with white, peristomal callus. Peristome: sharp, always inter- 
rupted for a short distance on the parietal wall; well and 
rather abruptly expanded along the palatal and basal walls, 


50 University of Michigan 


lobate at parietal angle and quite markedly auriculate at basal 
angle; but very slightly expanded in columellar region; thick- 
ened internally by continuous callus, which, however, is thin 
in the parietal and very narrow in the columellar region. 
Operculum: 314 whorls; similar to that of T. aurantia, but 
with caleareous portion much thinner; calcareous plate closely 
applied to horny base; externally the nuclear concavity and 
the linear convexity are much less prominent than in aurantia. 
Radula: similar to muskusi. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam. min.diam. alt. diam. long wide 


male 10.5 60(6.3) 47(4.9) 45(4.7) 92(4.3) 35(3.7) 73(2.7) 4% 
female 
(type) 14.0 54(7.6) 44(6.2) 44(6.2) 84(5.2) 34(4.7) 77(3.6) 5% 


Tudora rupis newportensis, new subspecies 

Type locality: (C1) along road, just north of New Port, 
Curacao. 

Distribution: Curacao; probably occurs on the more recent 
limestone all along the west shore, south of Spaansche Baai. 
Very common; on trees and brush and under rocks. 195 
specimens collected. 

Shell (fig. xii-D): considerably larger but relatively thin- 
ner than rupis. Color: like rupis, but spiral bands, although 
sometimes absent, usually are more distinct and more com- 
monly continuous; almost always with at least one continuous 
band surrounding the umbilicus from just below the parietal 
angle of the aperture. Whorls: usually more than 434 
retained. Last whorl (fig. x-30): growth-threads regular, 
closely-spaced, low and rounded, often so broad and coalescent 
as to render the surface practically smooth; spiral sculpture 
typically obsolescent except in umbilical region, where it de- 
velops low, rounded ridges, always surmounted by the growth 
sculpture. Earlier whorls: growth sculpture closely resem- 
bling that of awrantia; spiral sculpture very obscure. Peristo- 
mal callus: thinner than in rupis. Opereulum: 314 whorls. 
Other characters as in rupis. 


Occasional Papers of the Museum of Zoology 51 


Measurements 


Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 

male 
(type) 14.0 51(7.2) 44(6.1) 41(5.7) 88(5.0) 31(4.4) 80(3.5) 
female 15.8 58(9.2) 46(7.3) 43(6.8) 94(6.4) 34(5.3) 79(4.2) 
female 17.2 57(9.8) 45(7.8) 43(7.4) 87(6.4) 32(5.5) 80(4.4) 


or or ol 


Tudora muskusi muskusit H. Burrington Baker 


(1924; Naut., XX XVII, 93) 
?T. costata Vernhout (1914; Notes Leyden Mus., 180). 


Type locality: (C17b) top of shore cliffs near Knip Baai, 
northern Curacao. 

Distribution of species: Curacao; along western side, from 
north side of Plaja Abau to west side of entrance to Salinja 
Sint Marie. 1,252 specimens collected. 

Distribution of subspecies: Curacao; along western side, 
from both sides of Plaja Abau to Sint Jan Baai (C17, 16, 15), 
south of which subspecies grandiensis predominates. Quite 
abundant on trees and brush and under rocks; only found on 
or very near limestone. 881 specimens collected. 

Shell (fig. xii-C) : heavier, but slightly smaller, than awran- 
tia. Ground color: flesh-color to opaque whitish; spiral bands 
usually absent although single chocolate band not infrequently 
circles the umbilicus from the parietal angle of the aperture; 
eroded apex and apical plug white to bluish drab, usually very 
dark. Whorls: about 814, of which 5 are usually retained; 
not markedly convex, and with undulating suture rather well 
impressed. Last whorl (fig. x-31): growth costae quite 
regular but widely spaced, very prominent, heavy and angular, 
with a few, irregular, obsolescent cords between them; spiral 
sculpture of obsolescent, rounded thickenings, which are 
usually evident only as undulating buttresses along the sides 
of the growth costae ; towards the umbilicus the spirals become 
much more prominent, and surmount and break the growth 
sculpture into heavy bosses. Earlier whorls: growth sculpture 
lighter and broken by the prominent spiral sculpture into 


- 


52 University of Michigan 


bosses; similar to that of the last whorl of grandiensis. Sub- 
apical whorls (young shells) : similar to those of rupis but with 
heavier growth-costae. Umbilicus: rimate, even smaller than 
in rupts. Aperture: small, almost circular; with long axis 
inclined at about 45° to that of shell; internally light orange 
to chocolate-brown, with white, peristomal callus. Peristome: 
sharp, interrupted on parietal wall; not expanded at parietal 
angle, and with palatal wall slightly emarginate immediately 
below it; slight expansion of lower palatal and basal regions 
terminated abruptly just beyond slightly auriculate basal 
angle; columellar wall scarcely expanded; thickened internally 
by continuous, rounded callus, which is very narrow in the 
columellar region. 

Opereulum (fig. xi-43): 314 whorls; very similar to that 
of T. rupis. 

Radula (fig. xi-41): C/1; L1/1 +. 1/5-6; M/45-47. Inner 
lateral slightly more attenuate distad and spatulate cusps of 
outer lateral less variable than in 7. megachetlos; other char- 
acters very similar. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 12.4 56(7.0) 46(5.7) 40(5.0) 90(4.5) 30(3.7) 81(3.0) 5 
female 

(type) 15.8 56(8.9) 48(7.6) 40(6.3) 89(5.6) 34(5.3) 79(4.2) 5 
female 16.4 55(9.0) 48(7.9) 38(6.2) 97(6.0) 31(5.1) 78(4.0) 5% 
female 17.0 53(9.0) 43(7.2) 37(6.3) 89(5.6) 31(5.3) 79(4.2) 6 
T. costata; 

fig. (1846— 

1849) 15.7 52(8.2) 44(6.9) 78(5.4) 6 


Cl7be. Quite common on both escarpments of Plaja Abau; very com- 
mon near Seroe Djerimi, between Knip Baai and Plaja Djerimi (165 
specimens collected). Type lot (see Table V). 

Cl7a. Quite abundant; this lot is from the top of Seroe Djerimi (128 
specimens). Somewhat smaller, and with a slightly more noticeable 
tendency for the spiral sculpture of the last whorl to crenulate the 
growth costae. 

C16. Quite abundant (157 specimens). Color and shape similar to 
typical lot, but would average a trifle larger. Growth costae noticeabiy 


Occasional Papers of the Museum of Zoology 53 


lower, sharper and more compressed, but widely spaced. Spiral sculpture 
as weak as in typical lot. 

C15. Quite abundant; this lot was accidentally mixed with that from 
the eastern escarpment of Seroe Djerimi (Cl7a and C15; 431 specimens). 
By comparison of this mixed set with the other lot from Seroe Djerimi, 
it is very evident that the shells from C15 are similar to those from C16, 
but show still finer and more closely-spaced growth-costae. In addition, 
the spiral sculpture tends to approach that of the next subspecies. 


This species appears to be quite similar to TZ. costata 
(‘‘Menke’’ Pfr.) (1846; Zeit. Mal., 47) in color and sculpture ; 
the habitat of the latter is unknown. However, if the figure 
(1846-9; Chemn., II, fig. ix-9, 10) is even approximate, the 
much larger aperture of that species has its long axis almost 
parallel to that of the shell, and the parietal angle is even more 
markedly lobate than in 7. rupis. I do not believe that 7. 
costata occurs in the Dutch Leeward Islands, as none of the 
thousands of specimens examined by me shows any tendency 
to approach this striking aperture. Probably the specimens 
quoted by Vernhout (1914) belong to one of the subspecies of 
T. muskusi. 


Tudora muskusi grandiensis, new subspecies 


Type locality: (C14) northern half of Seroe Grandi, south 
of Sint Jan Baai, northern Curacao. 

Distribution: Curacao; along western side, from Seroe 
Baha So (4 specimens), to (at least) Seroe Grandi (C15, 14) ; 
southern limits undetermined. 317 specimens collected. 

Shell (fig. xii-F): smaller and slightly thinner and more 
slender than muskust. Color: spiral bands absent or present, 
rarely several continuous bands, frequently broken spirals, but 
most commonly with blotches of color connected to form un- 
dulating bands parallel to the growth lines; eroded apex and 
apical plug usually whitish or light yellow, and rarely as dark 
as normal muskusi. Sculpture of last whorl (fig. x—32): 
growth-costae much lower, and more closely-spaced (appar- 
ently due to the development of the interstitial cords) than in 
muskusi; spiral sculpture typically almost as prominent on the 


54 University of Michigan 


sides as around the umbilicus; it breaks the growth sculpture 
into elliptical bosses, or even produces a cancellate appearance. 
Earlier whorls: markedly cancellate, with cusp-like bosses at 
the intersections. Umbilicus: a trifle larger than in muskusv. 
Peristomal callus: thinner than in muskust. Other characters 
as in muskust. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 11.0 56(6.1) 46(5.1) 41(4.5) 89(4.0) 31(3.4) 79(2.7) 4% 
male 11.6 54(6.2) 48(5.0) 39(4.5) 85(3.8) 33(3.8) 74(2.8) 5% 
female 14.0 57(8.0) 48(6.7) 39(5.5) 91(5.0) 34(4.8) 75(3.6) 5 
female 

(type) 14.9 57(8.5) 48(7.1) 40(6.0) 88(5.3) 34(5.0) 72(3.6) 5% 


C14. Very common (311 specimens collected). Typical lot (see Table 
V); includes many specimens with fine, rather closely-spaced growth- 
costae, cut by poorly-developed spiral sculpture. 

C15. Very rare. Amongst the large numbers of shells examined in the 
field, 6 beautifully cancellate specimens were separated as quite character- 
istic of grandiensis. One other shell gives the appearance of a hybrid 
between T. fossor djerimensis and T. muskusi; a similar specimen comes 
from Cl7a (fig. xii-H). 


This subspecies intergrades with the more southern forms 
of muskusi, but not with the typical form. 


Tudora muskusi bullenensis, new subspecies 


Type locality: (C13) top of Seroe Largoe, just southwest of 
entrance to Salinja Sint Marie, near Sint Willebrordus, north- 
ern Curacao. 

Distribution: only collected from the top of this hill; it was 
not found at the base of the escarpments, or on the hill east 
of the entrance to the salinja. Infrequent, on trees and brush 
and under rocks. 54 specimens collected. 

Shell (fig. xii-G) : thinner and more slender than grandi- 
ensis. Ground color: bleached; whitish to pinkish; spiral 
bands absent, except for a very faint umbilical band in a few 
specimens; eroded apex and apical plug yellowish, pinkish or 
purplish-drab ; dilute varices of these last colors also common. 


Occasional Papers of the Museum of Zoology 55 


Whorls: about 514 retained. Sculpture (fig. x-33): low 
and weak; closest to grandiensis but much more obscure ; sides 
of last whorl cancellate or malleate; even the umbilical sculp- 
ture low and rounded. Earlier whorls: rather weakly can- 
cellate. Other characters similar to grandiensis. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 12.2 54(6.6) 44(5.4) 39(4.7) 87(4.1) 30(3.7) 70(2.6) 5% 
male 13.0 52(6.7) 42(5.4) 38(4.9) 94(4.6) 32(4.1) 76(3.1) 5% 
female 

(type) 16.4 50(8.2) 44(7.1) 37(6.0) 85(5.1) 28(4.6) 75(3.5) 6 
female 17.7 51(9.0) 42(7.5) 36(6.4) 88(5.6) 29(5.2) 79(4.1) 6% 


Section Tudora s. s. 


Shell: elongate to obovate conic; males average smaller than 
females, but the two sexes markedly intergrade in size. Sculp- 
ture: very variable; no definite tendency to accentuate the 
spiral ridges of the umbilical region. Post-embryonic whorls: 
color bands and spiral sculpture begin on 3rd whorl. Peri- 
stome: sharp, interrupted on parietal wall; more expanded 
in columellar than in palatal and basal regions; parietal angle 
produced and adjacent upper palatal wall emarginate. Oper- 
culum: calcareous plate well developed, practically equal in 
area to horny base; supporting lamellae high, thickened, and 
usually confluent; outer edge of caleareous portion only shal- 
lowly suleate. Radula: see megacheilos. 

Monotype: Cyclostoma simile ‘‘Gray’’ Sowerby. This sec- 
tion appears to be restricted to the islands of Aruba and 
Curacao. 


Tudora megacheilos megacheilos (Potiez and Michaud) 


Cyclostoma megacheilos Potiez and Michaud (1838; Gal. Douai, I, p. 
237, fig. xxiv-9, 10). C. simile ‘‘Gray’’ Sowerby (1843; Thes., fig. xxiv— 
48, 49). OC. megacheilum Pfeiffer (1846, Zeit. Mal., 33); first Curacao 
record. C. megachilum and megacheilus Pfr. (1847; Chemn., II, p. 66, 
fig. ix-18, 19). Tudora similis Gray (1850; Cat. Cycloph., 48) ; monotype 
of Tudora. Cyclostoma roridwm ‘‘Parr.’’ Pfr. (1852; Mon. Pneum. Viv., 


56 University of Michigan 


244); in synonymy. C. proteus ‘‘Beck’’ Pfr. (l.c.); in synonymy. 
C. cancellatum ‘‘Menke’’ Pfr. (l.c¢.); very large, light-colored female 
(cf. 1847; Chemn., II, fig. ix-15, 16, 17). 


Type locality: unknown; probably the Schaarlo, back of 
Willemstad, Curacao (C5c). 

Distribution of species: Curacao; western side, from 
Spaansche Baai to Kaap Sint Marie; eastern side from south 
of Sint Joris Baai to north of Landhuis Hato (at least) ; nar- 
rowly invades central region of older rocks. 1,911 specimens 
collected. 

Distribution of subspecies: Curacao; western side from 
Sint Anna Baai to Caracas Baai (C4, 5); intergrades with 
other subspecies to north and south. Under limestone rocks 
and on trees and brush; subarboreal, but not so much so as 
the species of Tudorata; does not penetrate the region of the 
older rocks to any great extent. 714 specimens collected. 

Shell (fig. xiii-A.) : subacuminate (with concave lateral out- 
lines) ; obovoid-conic; quite solid. Ground color: whitish to 
pinkish and dull amber; spiral bands present or absent, con- 
tinuous or broken; blotches rarely unite to form irregular 
bands parallel to growth lines; eroded apex and apical plug 
from almost white to orange-red and purplish black.** Whorls: 
about 714, of which 4 are usually retained; quite convexly 
rounded and with well marked suture; last whorls increase 
more rapidly in diameter than the earlier ones. Sculpture of 
last whorl: growth threads regular, closely spaced to con- 
tinuous, low and rounded; spiral sculpture present (typical 
megacheilos) to absent (form desculpta new; fig. xiii-B; 
x—34) ; when present it consists of few to numerous, low, 
rounded thickenings. Earlier whorls: similar to last whorl. 
Subapical whorls (young shells): almost as slender as in 
T. aurantia (1.e., with similar apical angle) ; with well-devel- 
oped, widely-spaced, growth lamellae and low, angular, spiral 
ridges. Umbilicus: of medium size, but almost hidden by 
peristome. Aperture: large, ovoid, with long axis at about 


38 1923; Occ. Papers Mus. Zool. Univ. Mich., no. 137, p. 3. 


Occasional Papers of the Museum of Zoology 57 


30° to that of shell; noticeably flattened on parietal wall; 
internally light buff to dark chocolate, usually showing the 
spiral bands when these are present; white peristomal callus 
often bordered internally by a dark band. Peristome: grad- 
ually but deeply expanded in palatal and basal portions, and 
broadly reflected, almost auriculate, in columellar region; 
parietal angle markedly produced; palatal emargination 
slight; internal callus quite thin, but continuous. 

Operculum :°° about 4 whorls; calcareous pla*e not exten- 
sively eroded over nucleus, elsewhere heavy, complete and dis- 
tant from the horny base; inner surface (horny plate) almost 
flat; outer surface (caleareous plate) concave at nucleus and 
with rim of each whorl slightly raised. 

Radula (fig. ix-27); C/1; L1/1+1/5-8; M/56-58. Cen- 
tral: shaped like a truncated, isosceles triangle; with very 
heavy base; single cusp stout and heavy; transverse thicken- 
ing a short distance below cusp. Inner lateral: similar to 
central but asymmetrical, with inner side longer and heavier 
than the outer; cusp longer and somewhat more slender. 
Outer lateral: long, rectangular base, with strong distal reflec- 
tion, which is pectinated into 5, spatulate, rather indefinite 
cusps (although 5 is the usual number, interstitial lobes or 
points may raise the number as high as 8) ; transverse thick- 
ening prominent, so as to produce a marked angulation on 
the sides of the base (as seen in profile). Marginal: with 
numerous, recurved, sharp cusps, which decrease in size 
towards the outer edge; the edges of the outer ones pectinate 
the distal edge of the tooth to fully 1/3 its depth; the un- 
divided, slightly thickened, recurved, outer portion slopes 
down abruptly to the comparatively narrow base. 


Measurements 
Shell Aperture Operculum Whorls 
alt. min.diam. alt. diam. long wide 
megacheilos P. and M. 
from text 
(1838) 15-18 67(10-12) 


39 L.¢., fig. ii-10. 


58 Unversity of Michigan 


Measurements (continued) 


Shell Aperture Opereulum Whorls 
alt. min. diam. alt. diam. long wide 
from figures 
(1838) 16.3 57( 9.3) 53(8.6) 90(7.7) 5 
simile Sowerby 
from figures 
(1843) 14.7 68(10.0) 48(7.0) 91(6.4) 414, 
19.0 66(12.5) 50(9.5) 85(8.1) 414 


cancellatum Pfr. 
from figures 

(1847) 20.3 69(14.0) 52(10.6) 80(8.5) 36(7.4) 68(5.0) 4% 
form desculpta, new 
male (C5cd) 13.1 69( 9.0) 55(7.2) 90(6.5) 37(4.9) 78(3.8) 3% 
female 

(type, Cded) 16.8 69(11.5) 52(8.8) 89(7.8) 37(6.2) 82(5.1) 4% 


Cded. Quite abundant (359 specimens collected). The Schaarlo (C5c) 
back of Willemstad is probably the type locality of this species, and this 
lot is that described above (see Table VI). It contains a large propor- 
tion of shells with vestigial spiral sculpture (form desculpta). 

C5ab. Quite abundant (186 specimens). Very similar to typical lot, 
but with considerably less tendency to approach form desculpta. 

C4. Quite abundant (169 specimens). Sculpture heavy, and with a 
tendency for the growth threads to coalesce. A few specimens are some- 
what cancellate, and thus show a slight tendency to approach kabrietensis. 
Form desculpta appears to be absent. 


Tudora megacheilos spreitensis, new subspecies 


Type locality: (C8) at base of northern escarpment of 
Seroe Spreit, Curacao. 

Distribution : Curacao; western side, from Kaap Sint Marie 
to Seroe Quinta, west to Sint Anna Baai, where it inter- 
grades with typical megacheilos and form desculpta; invades 
the region of the older rocks at Campo Bleinheim, and prob- 
ably intergrades with rondeklipensis east of Bullen Baai (C6, 
7, 8, 9,13, Ch6b). 548 specimens collected. 

Shell (fig. xiii-C): considerably heavier and larger than 
typical megachetlos. Color: similar in pattern to megacheilos, 
but usually more vividly colored and more distinctly banded. 
Seulpture of last whorl (fig. x-35): growth-cords heavier, 
more prominent, and more widely spaced than in megacheilos; 


Occasional Papers of the Museum of Zoology 


59 


spiral sculpture always present; higher but more compressed 
than in megacheilos; surmounted by growth sculpture which 
is developed into heavy, elliptical bosses; the last give the 
Earlier whorls: distinctly can- 


shell a granulate appearance. 


cellate, but without definite bosses or cusps at the intersec- 
tions. Peristomal callus: heavier than in megacheilos. 
characters as in megacheilos. 


Table VI. Dimensions of Tudor megacheilos and T. pilsbryt 


Other 


spreit-  spreit- rondeklip- mega-  kabriet- 
Subspecies ensis ensis ensis cheilos ensis pilsbryi 
Locality C13 C8 Cl2a Cded C3 C2ab 
Number of individuals 16 65 32 77 21 63 
27 98 83 158 23 132 
Mean number of 
whorls remaining 3.9 4.0 3.9 3.9 4.0 3.5 
4.1 4.3 4.1 4.1 4.3 3.6 
Extremes, ditto 3.34.0 354.5 3543 3545 3545 3.0-4.0 
4.0-5.0 3848 3545 3565 3848 3.0-43 
Mean altitude 14.8 14.0 13.7 13.2 13.9 13.0 
ae 16.4 16.5 15.5 15.8 14.7 
Mean index (1d/a) 56 54 53 54 54 64 
55 53 53 54 53 64 
Mean lesser diameter 8.2 7.5 7.3 Tal 7.4 8.4 
9.4 8.7 8.7 8.4 8.4 9.3 
Extremes, ditto 7.6— 8.6 6.7-8.0 65- 7.7 63- 7.5 7.0-7.8  7.7—- 8.7 
8.7-10.1 8.1-9.6 7.8-10.4 7.6-10.6 8.0-9.1 8.8-10.5 
Mean altitude; 
4 whorls 15.0 13.9 13.9 13.3 14.0 13.8 
17.0 15.7 16.0 15.3 15.6 15.3 
Extremes, ditto; 
4 whorls 14.0-15.6 11.5-15.2 12.7-14.8 12.2-14.3 13.1-14.7 12.2-15.8 
16.1-17.8 14.2-17.4 14.2-19.0 13.4-18.7 15.1-16.3 13.5_-17.6 
Mean index;4whorls 54 54 53 54 54 61 
55 55 53 55 56 62 
Extremes, ditto; 
4 whorls 53-56 51-58 51-55 51-56 53-55 54-65 
54-57 52-58 50-59 51-61 54-56 54-67 
Measurements 
Shell Aperture Opereulum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 14.9 67( 9.9) 54(8.0) 51(7.6) 96(7.3) 37(5.5) 80(4.4) 4 


female 


(type) 18.3 67(12.3) 53(9.6) 50(9.2) 100(9.2) 34(6.3) 78(4.9) 434 


60 Unversity of Michigan 


C8. Quite abundant (163 specimens collected). Type lot (see 
Table VI). 

C13. Common (43 specimens). Heavier; a larger proportion of can- 
cellate specimens. Although Tudora fdssor djerimensis also occurs in 
this locality, the two forms show no tendency to intergrade; in fact, the 
distinguishing characteristics appear to be actually accentuated (see 
tables VI and VII). 

C9. Infrequent (4 specimens). 

C7ab. Quite abundant (144 specimens). Very typical spreitensis. 

C6a. Very common (110 specimens). Bleached, more depressed and 
with sculpture weaker. Quite a number of specimens approach form 
desculpta. 

Cbh6b. Frequent (79 specimens). Similar to C6a, but even more 
bleached. 


Tudora megacheilos rondeklipensis, new subspecies 


Type locality: (C12a) base of northwestern escarpment of 
Ronde Klip, eastern Curacao. 

Distribution: Curacao; typical form only from the top and 
sides of Ronde Klip (C12); but the shells along the eastern 
limestone rim, from south of Sint Joris Baai to north of Campo 
Hato (C10, 11), are closer to this subspecies than to the others, 
although numerous specimens approach form desculpta; nar- 
rowly invades the region of the older rocks (Cb10). Habitat 
similar to megachetlos. 585 specimens collected. 

Shell (fig. xii—D): slightly larger and heavier than mega- 
cheilos. Color: similar in pattern to megacheilos, but the 
ground color is brighter and more pronounced, and the bands 
are usually obscure. Sculpture of last whorl (fig. x—36) : 
growth cords similar to spreitensis but finer and more com- 
pressed; spiral sculpture of numerous, prominent and sub- 
angulate ridges, which are scarcely surmounted by the growth 
sculpture and without bosses or distinct cusps at the intersee- 
tions. Earlier whorls: somewhat cancellate, but with the 
spirals much heavier than the growth sculpture. Umbilicus: 
more open than in megacheilos. Peristome: similar to, but 
more extensively expanded than that of megacheilos; inner 
callus heavier. Other characters as in megacheilos. 


Occasional Papers of the Museum of Zoology 61 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 
male 14.6 68( 9.9) 53( 7.7) 51(7.4) 95(7.0) 32(4.7) 83(3.9) 4% 
female 
(type) 19.0 68(13.0) 53(10.1) 52(9.9) 96(9.5) 33(6.2) 79(4.9) 4% 


Cl2a. Very common (115 specimens collected). Typical lot (see 
Table VI); on account of the great expansion of the peristome, the 
dimensions give the impression of a more elongate shell than is actually 
the case. 

C12b. Common (89 specimens). Typical rondeklipensis, but more 
bleached in color. 

Cllab. Very common (11b) to frequent (lla); 108 specimens. 
Similar to 12b, but with sculpture considerably reduced. A few closely 
approach form desculpta. 

Clied. Quite abundant (176 specimens). Bleached and rather small. 
A very large proportion approach form desculpta, and in several speci- 
mens the spiral sculpture is absent on the last whorl. 

C10. Infrequent; living specimens mainly on Cereus and under rocks 
at the base of these cacti (95 specimens). Big, heavy shells, usually with 
well developed sculpture, although a few show an approach to desculpta. 
Very much bleached in color and stained with a rusty deposit; even more 
so than Tudora aurantia wassauensis from B9, where the conditions are 
somewhat similar. 

Cb10. Rare (2 specimens). 


Tudora megacheilos kabrietensis, new subspecies 


Type locality: (C3) low, limestone escarpment near shore 
of Caracas Baai, and a short distance north of Fort Beeken- 
burg, southern Curaeao. 

Distribution: Curacao; near western shore, from New Port 
to Caracas Baai (C1, 3), north of which it intergrades slightly 
with typical megacheilos. 69 specimens collected. 

Color: similar to megachetilos, but bleached; pattern ob- 
secure. Sculpture of last whorl (figs. xiii-E) : numerous, com- 
pressed, angular, spiral ridges, surmounted by the growth 
sculpture, which develops slight, subpyramidal cusps at the 
intersections. Interior of aperture: white to dark orange and 
light chocolate; usually with markedly darker, spiral bands. 
Peristome: more deeply and extensively expanded than in 
megacheilos. Other characters similar to typical subspecies. 


62 University of Michigan 


Measurements 


Shell Aperture Opereulum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 
(type) 13.9 72(10.0) 54(7.5) 57(7.9) 94(7.4) 34(4.7) 79(3.7) 4 
female 16.5 66(10.8) 54(8.9) 49(8.0) 96(7.7) 32(5.3) 79(4.2) 41% 


C3. Frequent (44 specimens). Type lot (see Table VI); bleached 
and dull-colored. 

Cl. Infrequent (25 specimens). Lighter ground color and brighter 
spiral bands. This lot does not appear to approach T. pilsbryi any 
closer than does that from C3. 


Tudora pilsbryi H. Burrington Baker 
(1924; Naut., XX XVII, 94) 


Type locality: (C2a) at base of northern cliffs of the Tafel- 
berg of Santa Barbara, southern Curacao. 

Distribution: Cura¢ao; only collected near the base of the 
escarpments on the northern and western sides of the Tafel- 
berg (C2). Quite common; under limestone rocks, in crevices 
of the talus, and buried deeply in the soil; a few on the trunks 
of the larger trees; a distinctly terrestrial species. 195 speci- 
mens collected. 

Shell (fig. xiii-F): subacuminate, obovate-conic; much 
thinner and more depressed than megacheilos, but extremely 
variable in dimensions. Color: vivid lemon-yellow to bright 
salmon and dark plum-colored; usually uniform, but some 
of the lighter shells show darker, spiral bands of variable 
width, while others show darker varices; sculpture white; 
eroded apex and apical plug usually much darker, but of 
similar color to the remainder of the shell. Whorls: about 
7, of which 314 are usually retained; very convexly rounded 
and with very deeply impressed sutures; last whorls increas- 
ing more rapidly in diameter than the subacuminate, earlier 
ones, although, as in many species with a scalariform tendency, 
this character is very variable. Sculpture of last whorl (fig. 
x-39): regular, compressed growth-riblets, which are quite 
closely spaced but are narrower than their interspaces; spiral 


Occasional Papers of the Museum of Zoology 63 


sculpture of numerous (19 to 28 counted), regular, compressed, 
angular ridges, which are more widely spaced on the base than 
at the sides of the whorl, and heavier and slightly higher than 
the growth-riblets, which surmount them; intersections de- 
veloped as very prominent, sharp, pyramidal cusps. Earlier 
whorls: spiral and growth sculpture weaker and about equal; 
distinctly cancellate; cusps at intersections finer, but high 
and sharp. Subapical whorls: sculpture and shape similar to 
megacheilos (i.e., apical angle similar to aurantia). Umbili- 
cus: much larger than in megacheilos, and but little hidden by 
the peristome. Aperture: inner outline almost circular; in- 
ternally light buff to dark chocolate, usually with darker, 
spiral bands. Peristome: roughly triangular and markedly 
undulate; parietal angle produced and lobate; palatal emargi- 
nation a marked sinus; lower palatal wall extensively devel- 
oped but scarcely expanded; basal wall abruptly expanded; 
basal angle flattened; columellar wall auriculate below, but 
narrowed abruptly a short distance from the parietal wall; 
internal callus very poorly developed, practically absent on 
parietal wall. 

Opereulum (fig. xi44): about 4 whorls; similar to mega- 
cheilos, but with inner surface (thin, chondroid plate) slightly 
convex, and outer surface (caleareous plate) deeply and quite 
regularly concave over its entire surface. 

Radula: similar to megacheilos. 


Measurements 


Shell Aperture Operculum Whorls 
alt. maj.diam. min.diam. alt. diam. long wide 
male 13.2 85(11.2) 64( 8.5) 61(8.1) 93(7.5) 41(54) 80(4.3) 334 
female 
(type) 16.0 78(12.4) 63(10.1) 57(9.1) 98(8.9) 39(6.2) 81(5.0) 334 


Tudora fossor fossor H. Burrington Baker 
(1924; Naut., XX XVII, 94) 


Type locality: (Cb17b) valley between Seroes Palomba and 
Baha Hoendoe, northern Curacao. 


64 University of Michigan 


Distribution of species: Curaéao; western side north of 
Bullen Baai, and east in the northern hill to (at least) the 
Tafelberg of Sint Hyronimus. A single, dead specimen (acci- 
dental?) from Seroe Spreit. Aruba: all of the limestone por- 
tion and narrowly invades the metamorphic rocks. Almost 
entirely terrestrial in habits. 1,460 specimens collected. 

Distribution of subspecies: Curacao; north of Landhuis of 
Campo Sint Kruis; mainly restricted to wooded valleys and 
rocky hillsides in the central region of older rocks (Cb16, 
17abe; 20); also a somewhat different form from the Tafel- 
berg of Sint Hyronimus (C20). Frequent in well-wooded 
valleys; aestivates deep down in the talus and rock crevices; 
comes to the surface during rains but appears never to go far 
from the ground, even on the trunks of the larger trees. 282 
specimens collected. 3 

Shell (fig. xiii-G) : elongate-conic; rather thin. Color: last 
whorl dark plum-colored (dark, smoky amber by transmitted 
light), shading on the penultimate whorl into the dark horn 
color of the earlier whorls, which show indefinite, dark, spiral 
bands; sculpture tipped with bluish white; eroded apex and 
apical plug dark horn color to orange and plum color. Whorls: 
6 (in one, entire, male shell), of which about 31% are retained ; 
more convexly rounded and with deeper suture than in mega- 
cheilos; the diameter increases gradually and regularly from 
the first to the last. Sculpture ofi last) whorl (fig. x-37) : 
similar to pilsbryi; regular, compressed growth-riblets, which 
are closely spaced but much narrower than their interspaces; 
spiral sculpture of numerous (15 to 21 counted), regular, 
compressed ridges, which are more widely spaced in the umbili- 
cal region than on the sides, and are considerably heavier than 
the growth-riblets but are surmounted by them; intersections 
developed as lanceolate cusps. Earlier whorls: similar to 
pilsbryi; cusps at intersections less pronounced. Subapical 
whorls (young shells) : sculpture similar to those of megachet- 
los; not so slender (i.e., apical angle greater). Umbilicus: 
smaller than in megacheilos; partially hidden by peristome. 
Aperture: relatively smaller than in megacheilos, but similar 


Occasional Papers of the Museum of Zoology 65 


in shape; internally light to dark chocolate, when light it 
shows darker, spiral bands; peristomal callus white. Peri- 
stome: palatal and basal walls rather narrowly but quite 
abruptly expanded; columellar region more extensively ex- 
panded (although less so than in megacheilos) ; parietal angle 
produced; palatal emargination very slight (fig. xili-K) ; in- 
ternal callus extensive but very thin, practically absent on 
parietal wall. 

Operculum: 314 whorls; similar to megacheilos, but nuclear 
erosion more extensive, caleareous portion slightly smaller than 
chondroid plate, and entire outer surface (calcareous plate) 
distinctly concave, although much less so than in T. pilsbryi. 

Radula: similar to megachetlos. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 
male 
(type) 13.0 58(7.5) 48(6.2) 45(5.9) 93(5.5) 37(4.8) 79(3.8) 6(en- 


tire) 

female 15.0 61(9.2) 49(7.3) 48(7.2) 93(6.7) 35(5.3) 76(4.0) 4 

Cb17b. Frequent (195 specimens collected). Typical lot (see Table 
VII) ; coloration very uniform. . 

Cb17a. Apparently infrequent, but the abundance of this sub-species 
can only be judged during rain-storms (13 specimens). Typical fossor. 

Cb16. Very rare; 1 dead, bleached specimen. 

Cb20. Apparently quite rare (3 specimens). Typical fossor reaches 
an altitude of around 1,200 feet on Sint Christoffelberg. 

C20. Frequent (70 specimens). Slightly larger, with lighter ground- 
color, and much more variegated color; a quite divergent form. 


Tudora fossor djerimensis, new subspecies 


Type locality: (C17b) top of shore cliffs between Knip Baai 
and Plaja Djerimi, northern Curacao. 

Distribution: Curacao; western limestone rim from both 
sides of Plaja Abau to Salinja Sint Marie (C138, 14, 15, 16, 
17); also one, dead, bleached specimen from Seroe Spreit 
(C8). Quite infrequent to very rare; under limestone rocks; 
almost completely terrestrial. 207 specimens collected. 


66 University of Michigan 


Shell (figs. xii-I) : slightly larger and considerably heavier 
than typical fossor. Ground color: bleached; white to light 
pinkish ; spiral bands absent or present, entire or with blotches 
united to form bands parallel to the growth line; eroded apex 
and apical plug light buff, to orange and deep indigo. Whaorls: 
about 384 retained. Sculpture: similar to fossor throughout, 
but heavier and with comparatively less prominent cusps; 
spiral ridges usually less crowded on sides of last whorl. Aper- 
ture: internally light orange to light chocolate; usually darker, 
spiral bands can be seen. Peristome: more extensively but 
less abruptly expanded than in fossor; internal callus much 
heavier, continuous. Other characters as in fossor. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 13.7 63( 8.6) 50(6.8) 48(6.6) 88(5.8) 35(4.8) 79(3.8) 334 
female 

(type) 18.0 62(11.2) 48(8.7) 45(8.0) 93(7.4) 32(5.8) 76(4.3) 41% 
Cs 

(male) 14.7 60( 8.8) 48(7.0) 47(6.9) 91(6.3) lacking 414 

Cl17be. Quite infrequent (58 specimens collected). Type lot (see 
Chart VII). 

Cl7a. Quite infrequent, on top of Seroe Djerimi (42 specimens). 
Typical. 

C16. Infrequent (23 specimens). A little larger than the typical lot. 

C15. Infrequent; accidentally mixed with a lot from the eastern es- 
carpment of Seroe Djerimi (C15 and Cl7a; 57 specimens). Like C16. 

C14. Very infrequent (17 specimens). Rather large. 

C13. Quite rare (10 specimens). Smaller, but otherwise typical. 


Tudora fossor westpuntensis, new subspecies 


Type locality: (C18) small limestone remnant, less than a 
kilometer north of Plaja Abau, northern Curacao. 

Distribution: Curacao; limestone rim north of Plaja Abau 
(C18, 19). Under limestone rocks, and, during rains, on bases 
of vegetation; almost completely terrestrial. 252 specimens 
collected. 

Shell (fig. xiii-H): females about as large as males of 
fossor. Ground color: light buff to pink, orange and light 


Occasional Papers of the Museum of Zoology 67 


plum-color; blotches of broken spiral bands, when present, 
usually united to form irregular bands parallel to the growth 
lines; eroded apex and apical plug buff to orange and deep 
indigo. Growth sculpture of last whorl: heavier, higher and 
more closely-spaced than in fossor ; whorl] distinctly cancellate ; 
cusps less prominent than in fossor. Whorls: about 314 re- 
tained. Umbilicus: less hidden by peristome than in fossor. 
Aperture: internally light buff to vivid orange; broken, spiral 
bands usually shown. Peristome: less abruptly expanded than 
in fossor. Other characters as in fossor. 


Measurements 


Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


male 
(type) 9.8 65(6.4) 52(5.1) 45(44) 93(4.1) 32(3.2) 80(2.5) 3% 
female 13.5 62(8.3) 49(6.6) 46(6.2) 92(5.7) 34(4.6) 78(3.6) 4 


C18. Frequent (145 specimens collected). Type lot (see Table VIZ) ; 
these smallest specimens, peculiarly enough, come from a habitat where 
Cerion wa attains greater than average size. 

C19. Infrequent; under limestone slabs at the very edge of the shore 
cliffs (107 specimens). Slightly larger; bleached and stained with a 
dark brownish deposit. 


Table VII. Dimensions of Tudora fossor40 


canashit- westpunt- djerim- 
Subspecies arubana ensis ensis fossor ensis 
Locality A2ab A4ab C18 Cb17b Cl17be 
Number of 
individuals 80 38 68 76 13 
120 64 ra 119 45 
Mean number of 
whorls retained 3.5 3.4 3.4 3.5 3.6 
3.7 3.5 3.6 3.6 3.9 


Extremes, ditto 3.04.0 3.0-6.0 3.0-3.8 3.0-6.0 3.3-4.3 
3.3-4.3 3.0-5.5 3.0-4.5 3.34.3 3.3-4.5 
Mean altitude 13.2 11.4 10.5 12.0 12.6 
15.7 13.4 12.4 14.1 15.0 


40 The upper horizontal row under each heading contains the data for 
the males; the lower, those for females. 


68 Umversity of Michigan 


Table VII.— (Continued) 
Dimensions of Tudora fossor 


canashit- westpunt- djerim- . 
Subspecies arubana ensis ensis fossor ENnsis 
Locality A2ab A4ab C18 Cbh17b C17be 
Mean min. diam. 
index 52 52 53 53 52 
52 52 52 52 51 
Mean min. diam. 6.9 5.9 5.5 6.3 6.6 
8.1 6.9 6.4 7.3 7.6 


Extremes, ditto 6.0—7.4 5.0-6.3 5.1-5.9 5.9-6.7 6.1-6.8 
7.5-8.8 6.4-7.6 6.0-7.1 6.8-8.1 6.9-8.8 


Mean altitude; 
3144 whorls 13.2 11.3 10.7 12.3 12.6 
15.3 13.3 12.2 13.7 14.0 


Extremes, ditto; 
31% whorls 11.8-14.4 10.6-11.9 9.6-11.6 11.4-13.1 11.6—13.7 
14.2-17.0 11.9-14.4 11.2-13.5 12.3-14.9 13.2-14.6 


Mean min. diam. 
index; 314 whorls 52 52 52 52 52 
53 52 52 52 52 


Extremes, ditto; 
31% whorls 49-56 50-54 50-54 49-55 50-54 
50-56 50-55 50-55 50-55 50-54 


Tudora fossor arubana, new subspecies 


Type locality: (A2b) spur of Seroe Pretoe, between Roois 
Spoki and Hundoe, southern Aruba. 

Distribution: Aruba; limestone deposits southeast of 
Spaansch Lagoen (Al, 2). Quite common; mainly under 
limestone rocks, but also on trunks and the larger branches 
of trees in well-wooded places (A2); more nearly arboreal 
than the other subspecies. 490 specimens collected. 

Shell (fig. xiii-K) : averages the largest in the species, but 
scarcely as heavy as djerimensis. Ground color: cream to 
salmon and orange, the reddish tints predominate; spiral 
bands present or absent, continuous or broken, often with 
blotches joined to form varices; eroded apex and apical plug 
buff to (usually) deep indigo. Seulpture of last whorl (fig. 
x—38): prominent to almost confluent, closely-spaced to con- 
tiguous, regular growth-threads, which are more rounded 


Occasional Papers of the Museum of Zoology 69 


than in the other subspecies; spiral sculpture of low, weak, 
rounded to subangular ridges, which are often obsolescent, 
and are scarcely surmounted by the growth threads; inter- 
sections without cusps. Earlier whorls: closely-spaced, growth 
sculpture obscures the weaker, spiral ridges. Subapical 
whorls: like fossor. Umbilicus: even less hidden by the 
peristome than in westpuntensis. Aperture: internally light 
orange to dark chocolate, usually vivid orange; spiral bands 
inconspicuous. Peristome: quite deeply but gradually ex- 
panded; internal callus quite heavy, continuous. Other char- 
acters as in fossor. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 


. male 13.0 65( 8.4) 52(6.7) 51(6.6) 91(6.0) 35(4.5) 73(3.3) 31% 
male 13.7 63( 8.6) 50(6.9) 48(6.5) 91(5.9) 36(4.9) 74(3.6) 344 
female 

(type) 16.2 62(10.0) 50(8.1) 47(7.6) 95(7.2) 33(5.3) 76(4.0) 4 


A2ab. Common (200 specimens collected). Type lot (see Table VII) ; 
many specimens have the spiral sculpture considerably reduced (fig. 
xiii-5), 

A2e. Quite common; under limestone slabs (161 specimens). Slightly 
larger, but otherwise typical. 

Al. Quite infrequent; under limestone slabs (129 specimens). 
Bleached; almost half of the shells uniform whitish to pinkish. Heavier, 
with more prominent sculpture, and with especially well-developed 
peristomal callus. 


This subspecies is, by itself, distinct enough to be a separate 
species, but the next form, which is plainly a geographical 
variety of it, is so close to westpuntensis that it was thought 
best to regard all of the Aruba specimens as subspecies of 
fossor. 


Tudora fossor canashitensis, new subspecies 


Type locality: (A4b) top of Seroe Canashito, Aruba. 

Distribution: Aruba; limestone rim from Spaansch Lagoen 
and Boedoei north; narrowly invades region of older rocks at 
the Hooiberg (A3-8). Under rocks; a distinctly terrestrial 
form. 229 specimens collected. 


70 Unversity of Michigan 


Shell (fig. xiii-I) : somewhat smaller than fossor. Ground 
color: buff to salmon; the reddish tints predominate; spiral 
bands absent or present; usually with the blotches of the 
broken bands united to form varices parallel to the growth 
lines; eroded apex and apical plug buff and light orange to 
(usually) dark plum-color. Whorls: about 314 retained. 
Seulpture of last whorl: growth-ribs closely spaced, and 
heavier than in fossor, but scarcely surmounting the spiral 
sculpture which is weaker than in fossor; intersections seldom 
developed into cusps. Umbilicus: like arubana. Peristome: 
columellar expansion weaker than in westpuntensis. Other 
characters as in westpuntensts. 


Measurements 
Shell Aperture Operculum Whorls 
alt. maj.diam.min.diam. alt. diam. long wide 
male 
(type) 12.5 57(7.1) 48(6.0) 44(5.5) 89(4.9) 32(4.0) 78(3.1) 6(all) 
female 14.4 58(8.3) 49(7.1) 45(6.5) 89(5.8) 33(4.8) 77(3.7) 3%4 


A4ab. Quite infrequent; more common on top than at the base of the 
escarpment (102 specimens collected). Type lot (see Table VII). 4 
A5. Infrequent; under limestone slabs near the edge of the shore cliffs 
(45 specimens). Slightly smaller. 

A6. Rare (10 specimens). Although dead shells are frequent through- 
out the plantations of aloes, living specimens are very rare. 

Ava. Rare (2 specimens). Bleached and stained. 

A7b. Quite rare (12 specimens). Bleached, and stained with a rusty 
deposit; very close to westpuntensis from C19. 

Avec. Very rare; 1 dead, bleached specimen. 

A3. Frequent; at the base of the escarpments (57 specimens). Simi- 
lar to those from A4b; but somewhat approaches arubana. 


AMNICOLIDAE 


Potamopyrgus parvulus (Guilding) 


Paludina parvula Guilding (1828; Zool. Jour., III, pp. 537-8; Suppl. 
Pl. xxviii, figs. 1-3). Paludina jamaicensis C. B. Adams (1849; Cont. 
Conch. 42). Paludestrina crystallina Smith (1898; Proc. Mal. Soe., III, 
113); first Bonaire record; collected by Hartert. 


Type locality: St. Vincent, Lesser Antilles. 


Occasional Papers of the Museum of Zoology - 71 


Distribution: Jamaica, Haiti, St. Thomas, St. Vincent, 
Grenada. Curacao: Sint Marie Spring (Ce13). Aruba: reser- 
voir and brook at Fontein (Ac2). Bonaire: well in Kralendijk 
(not collected), and Pos Baca (Bel). Klein-Bonaire: Pos di 
Cas (Kel). 

Guilding’s figures 1 and 2 show both the shell and the 
animal; figure 3 is apparently a young specimen. They all 
represent the smooth form, but show evident, spiral striations. 
From authentic specimens in the A. N. 8S. P. (no. 67466, from 
C. B. Adams), P. jamaicensis appears to be a synonym. This 
species is more slender and slightly smaller than P. coronatus 
(Pfr.) ; parvula is, of course, the older name. Although the 
original figure looks very much like parvulus, Paludestrina 
valenciae Preston (1909; Ann. Mag. Nat. Hist., III, p. 513, 
fig. 16), from Lake Valencia, Venezuela, is still smaller and 
more slender; its last whorl is shorter in proportion to the 
spire, and the aperture is more nearly circular (A. N. 8S. P., 
no. 99401, three cotypes). 

All of my specimens are dull greenish in color. Those from 
Curacao, Aruba and Bonaire belong to the typical, smooth 
form (figs. xi-45, 46), but the lot from Klein-Bonaire contains 
a small proportion of weakly spinose shells (fig. xi47). How- 
ever, none is as markedly angulate as P. coronata, and the 
black-margined thread-carina is simply erenulated into low, 
triangular cusps. In my specimens, this carina begins as a 
slightly raised line on the penultimate whorl, gradually de- 
velops into the crown of spines, and then dies out again 
towards the aperture. 


Measurements 
alt. maj. diam. alt. apert. diam. apert. whorls 
Figure 45 4.62 54(2.49) 37 (1.73) 78 (1.35) 6 
Figure 46 4.48 49 (2.19) 36 (1.63) 76(1.25) 64 
Figure 47 3.60 58 (2.10) 41(1.47) 77 (1.13) 5144 
PLANORBIDAE 


Planorbis pallidus C. B. Adams 


Planorbis pallidus C. B. Adams (1846; Proc. Boston Soc., II, 102). P. 
pallidus Clessin (1884; Chemn., II, p. 122, fig. xi-7). P. circumlineatus 


72 University of Michigan 


‘*Shuttleworth’’ Clessin (op. cit., p. 211, fig. xxxii-6) ; Sowerby (1878; 
Conch. Icon., XX, fig. vi-48). Planorbis sp.? Smith (1898; Proce .Mal. 
Soc., III, 113) ; first Curacao record of genus; collected by Hartert. 


Type locality: Jamaica. 

Distribution: Jamaica, Haiti, Porto Rico, St. Thomas. 
Curacao: cement tank in front of Landhuis Wilhelmina 
(Ce2); dammed pool on northwest side of Seroe Papaja 
(Cell; dead shells only); dammed pool, Campo Lagoen 
(Ce17). Bonaire: Pos Baca (Bel). The water in all of these 
places is slightly brackish to the taste, but it is used for stock 
animals; that from the first is quite drinkable. 


Measurements41 
Shell Aperture Whorls 

alt. maj. diam. min. diam. alt. diam. 
P. pallidus 
Adams (1846) 3.2 320(10.2) 3 (sic) 
Clessin (1884) 2.3 370( 8.5) + 
P. circumlineatus : 
Clessin 2.5 360( 9.0) 4-5 
P. weilandi 
Pfr. (1876) 2.5 320( 8.0) 220(5.5) (3.0) 4 
P. kiihnianus 
Clessin 2.0 325( 6.5) + 
P. meridaensis 
Preston 2.7 300( 8.0) 110(3.0) 1.5(sic) 3% 
Ce17 (largest) 3.0 355(10.6) 285(8.6) 120(3.6) 110(4.0) 5 

2.9 300( 8.7) 255(7.4) 120(3.5) 110(3.9) 4% 
Bel (largest) 2.6 360( 9.4) 310(8.1) 1380(3.4) 115(3.9) 4% 


Cel. Quite abundant (33 specimens collected). Thin, transparent and 
rather small. As the tank is artificial and its water supply comes from a 
well, the shells must have been introduced by some extraordinary means 
of dispersal. 

Cell. (6 dead shells collected). Opaque and rather heavy; similar 
to the next lot. 

Ce17. Infrequent (3 living and 15 dead shells). Large, rather heavy 


41 The altitude in my measurements is that opposite the aperture; from 
their figures, this seems to correspond most closely with that used by the 
older writers. The altitude given for P. meridaensis is estimated from 
the figure. 


Occasional Papers of the Museum of Zoology 73 


and stained with brownish. The whorls of this lot and the preceding one 
are somewhat higher and more swollen than in the other shells. 

Bel. Quite common (34 specimens collected). Thin, white and trans- 
parent; thickly covered with gelatinous material and usually with one 
to three egg-masses stuck to the shell. Mostly with one to four rather 
heavy white varices, which must correspond to resting periods in the 
growth of the shell; at these places, the plane in which the whorls are 
coiled is apt to change slightly but sharply, so that an irregular shape 
results. In the umbilical region of some specimens, the last whorl com- 
pletely covers a portion of the penultimate whorl. 


Description of Curacao specimens. Shell: dextral in form. 
Color: clear white and transparent, or opaque and stained 
with brownish; when alive it appears reddish with coppery 
shadows, as the color of the animal shows through. _Whorls: 
4 to 5; quite rapidly increasing in diameter; sutures shallow 
on umbilical side but deeper above. Last whorl: somewhat 
flattened above, less so below; rounded to scarcely subcarinate ; 
growth sculpture regular and well marked; impressed, spiral 
lines irregular and sometimes almost obsolete; not hispid. 
Earlier whorls: rounded above and below, but the succeeding 
whorls obscure the convexity in the umbilicus. Apical whorls: 
easily seen in umbilicus, but deeply sunken and scarcely visible 
from apical side. Umbilicus: shallowly and regularly concave ; 
less than 1/3 the major diameter of the shell. Aperture: 
oblique, very variable in shape; elliptical to semilunate; usu- 
ally twisted downwards, sometimes markedly so, but may even 
project slightly upwards. Peristome: simple, sharp; parietal 
eallus thin. 

P. pallidus has a characteristic tendency for each whorl to 
twist slightly downward from the plane of the preceding one; 
the amount of this is often irregular so that the umbilicus may 
be markedly elliptical and the exposed portions of the earlier 
whorls very variable in width. Adams described this species 
as scarcely 3-whorled, but he must have counted the whorls 
as visible from the apical side, as smaller specimens in the 
A. N. 8. P. (no. 62014, from Kingston, Jamaica, and other 
lots) show 4 whorls, are dull horn-colored, and more closely 


74 Umversity of Michigan 


approach Clessin’s description and figure. P. circwmlineatus 
Clessin is apparently a synonym, but Sowerby’s figure, 
although he also quotes Shuttleworth MSS., looks like a 
Planorbula. Specimens from St. Thomas (A. N.S. P., 123885) 
approach the Curacao shells in size, color and texture. In 
P. weinlandi Pfr. (1876; Mal. Bl., XXIII, p. 172, fig. ii-9, 10, 
11), from a brook near Jeremie, Haiti (A. N.S. P., 91455, one 
specimen from Weinland), the aperture encloses less of the 
penultimate whorl than in typical pallidus, but this character 
is very variable in the Curacao shells. P. meridaensis Preston 
(1907; Ann. Mag. Nat. Hist., XX, pp. 493, 497, fig. 18), from 
Merida, Venezuela, is a heavier shell with more swollen 
whorls, and with the umbilicus more deeply sunken and show- 
ing better impressed sutures (A. N. S. P., 98195; three 
cotypes). This last form is slightly larger than P. kiithnianus 
(+ kithnerianus) ‘‘Dunker’’ Clessin (op. cit., 108, fig. xi-12), 
from Surinam, but otherwise appears to satisfy the description 
and figure. All of these planorbes have sinistral shells, if the 
visibility of the first whorls is taken as the criterion of the 
apex. Otherwise, they appear to fall into the section 
Gyraulus, although the epidermis is not hispid. 


PUPILLIDAE*?4 


Gastrocopta longurio (Crosse) 


Pupa longurio Crosse (1872; J. de C., XX, 158); Crosse and Bland 
(1873; J. de C., XXI, fig. I-2). 


Type locality: ‘‘Curacao,’’ probably the Schaarlo (C5e), 
back of Willemstad. 

Distribution: Curacao, Aruba, Bonaire, Klein-Bonaire; 
almost everywhere under limestone rocks; apparently with 
preference for the more arid situations (C1—6, 10-13, 15, 17, 
18; A2-5, 8; B1-5, 8; K1). 

Numerous specimens of Gastrocopta and Pupoides were col- 
lected, and have been submitted to Dr. H. A. Pilsbry for study. 


41a See Addenda page 116. 


~l 


Occasional Papers of the Museum of Zoology 


The relationships and variation of the present species, and the 
identification of the next, will be discussed by him in a 
future paper. 


Pupoides sp? 

Pupa fallax Gibbons (1879; J. of C., II, 131); first Curacao record. 
Pupoides simoni? H. B. Baker (1923; this series, no. 137, p. 5). 

Distribution in the islands: Curacao, Aruba, Bonaire, Klein- 
Bonaire; with the preceding but not as abundant nor in quite 
as arid situations (C3—6, 12, 17; A3-5, 8; B2, 4,8; K1). This 
species is probably not limited to the islands, although it is 
omitted from the list in the discussion of zodgeographical 
affinities. 


SUCCINEIDAE 


Succinea gyrata Gibbons 
(1879; J. of Conch., IT, 136, fig. I-2) 


Type locality: ‘‘at St. Ann’s, Curacao’’; probably the 
Schaarlo (C5c), back of Willemstad. 

Distribution: Curacao (C5, 6, 11, 17; Cbh6, 17); Bonaire 
(B3; Bb5); Klein-Bonaire (K1). Rare to quite infrequent; 
under stones and the bark of dead trees, and in the crevices of 
the bark of living ones; abundance entirely independent of the 
nature of the underlying rocks. Most of the living specimens 
were juvenile or small, which leads me to believe that this 
species only attains maturity during the wet season. 62 
specimens collected. 

In what is here regarded as the type locality, the typical, 
elongate form of this species integrades with more globose 
specimens with relatively larger aperture, such as are repre- 
sented by the measurements of those from C11 (see below). 
The whorls are convex and the suture is deeply impressed, 
almost channeled. The surface has a peculiar, satiny luster, 
due to a microscopic sculpture of irregular, raised punctua- 
tions. The growth wrinkles are prominent and, towards the 
aperture, are often very pronounced, almost subcostate, and 


76 University of Michigan 


irregular. The shell is rendered imperforate by a slight, 
thickened reflection of the columella in the parietal region, 
where it is continued by a thin callus that connects the ends 
of the peristome. 


Measurements 
alt. maj.diam. alt.apert. diam.apert. whorls 
Gibbons (1879) 12.0 54(6.5) 58 (7.0) 64 (4.5) 4 
Cb6 (dead) 9.3 52 (4.8) 60(5.7) 65 (3.7) 314 
C11 (dead) 13.5 57 (7.7) 65 (8.8) 66 (5.8) 38% 
C11 (dead) 11.4 56 (6.4) 70(8.0) 64(5.1) 344 
Bbd (living) 9.3 57 (5.3) 65 (6.0) 67 (4.0) 3 


The radular formula (fig. xiv-48) of a medium-sized speci- 
men is 241-24. The central tooth is symmetrical and tri- 
euspid. The Ist to 7th laterals are asymmetrical but also tri- 
cuspid; the entocone is small, slightly below the level of the 
mesocone, and is obscured by the refraction of the edge of the 
tooth. All of the cusps are elongate and very sharp. The 8th 
lateral usually adds a minute, outer, 4th cusp. Beyond this, 
the number of cusps becomes very variable, as is illustrated 
by the three figured examples of the 14th tooth, which occur 
within four consecutive transverse rows. The 24th tooth is a 
mere denticle. 

The jaw (fig. xiv-49) of the same specimen is double, and: 
consists of a rounded-oblong, emarginate, basal plate and the 
recurved, arcuate true jaw.. The latter bears a triangular 
median thickening or low rib, and two, transverse, lateral ones; 
these slightly undulate both margins of the recurved portion. 
The outer surface is closely, but irregularly, striate; these 
markings converge towards the apex of the central triangle, 
and are parallel to the sides of the lateral thickenings. 


ZONITIDAE 
Guppya molengraaffi, new species 


Type locality: (Cb20) just west of the summit of Sint 
Christoffelberg, northern Curacao, at an altitude of about 
1,200 feet. 


Occasional Papers of the Museum of Zoology LE 


Distribution: Curacao; one specimen collected at type 
locality. | 

Shell (fig. xv-54): small, depressed turbinate; light horn- 
colored; thin and shining. Whorls: 414; convex; gradually 
and regularly increasing; suture shallow. Growth striae: well 
marked, especially on the base, which shows quite regularly 
spaced, impressed lines. Spiral striae: present, but much 
weaker and more irregular than is usual in the genus. Um- 
bilicus: small, about 1/12 of the major diameter of the shell; 
circular. Aperture: subvertical, reniform. Peristome: simple, 
sharp, incomplete. 

Radula: C/3; L5/3; M19/3 + 4/4-+- 1/1; or 25-5-1-5-23. 
Form of teeth very similar to those of G. gundlacht (Pfr.) ,* 
although the lateral cusps of the central are somewhat more 
acuminate and the outer cusp of the first marginal is smaller 
and more distant from the two larger ones. Jaw: much as 
in G. gundlachi (l.c., fig. 3). 


Measurements 
altitude maj. diam. min. diam. alt. apert. diam.apert. whorls 
1.63 150 (2.40) 130 (2.13) 63 (1.03) 120 (1.22) 44 


The practical absence of spiral sculpture makes this species 
appear closest to Guppya miamiensis Pilsbry (1903; Naut., 
XVII, 77) from Florida, but the Curacao shell is more globose, 
with a more depressed spire and shallower suture. The regu- 
larity of the impressed striations on the base is also a notice- 
able difference. 


HELICIDAE 


Thysanophora crinita arubana, new subspecies 


Type locality: (A4a) at base of northern escarpment of 
Seroe Canashito, Aruba. 


431922; Occ. Papers Mus. Zool. Univ. Mich.; no. 106, p. 45, fig. 
xvii-l. 


78 University of Michigan 


Distribution: Aruba; very rare; 6 specimens under lime- 
stone rocks at type locality. 

Shell (fig. xv-56): small, sub-discoid, rather heavy; dark 
horn-colored. Whorls: 414, markedly flattened above but 
seareely subangulate; gradually increasing; suture deeply im- 
pressed vertically ; last whorl slightly descending. Sculpture 
of last whorl: closely spaced, compressed, undulate and rarely 
anastomosing, cuticular riblets, which are slightly more ob- 
lique than (and cross) the obscured growth wrinkles. Penul- 
timate whorl: in addition, with the broken bases of what, in 
young specimens (fig. xv—55), are long (.33 mm.), sparsely 
and irregularly scattered, white hairs, which extend even into 
the umbilicus. Embryonic whorls: 114; with regular, some- 
what heavier, cuticular costae, which are slightly more oblique 
than the obscured growth wrinkles and extend to the very 
apex. Umbilicus: large, about 3/10 the major diameter of the 
shell; circular. Aperture: oblique, subcireular. Peristome: 
incomplete, very slightly thickened; parietal callus very weak. 


Measurements 
altitude maj. diam. min.diam. alt.apert. diam.apert. whorls 
2.09 190 (3.93) 160 (3.38) 61(1.27) 140 (1.76) 44 


These shells are certainly very closely related to Tri- 
chodiscina crinita Fulton (1917; P. Mal. Soc., XII, 240), 
from Carthagena, Colombia. However, the Aruba form is evi- 
dently a larger shell (even after making allowance for the 
greater number of whorls) ; its last whorl is markedly flattened 
above instead of evenly convex; the incomplete peristome is 
slightly thickened ; and the parietal callus is very weak, almost 
absent (from Fulton’s figure, it would appear that the peri- 
stome of his shell is continuous, but I suspect that it simply 
has a heavy, parietal callus). Both crinita and arubana must 
have similar sculpture to that in Trichia venezuelensis 
Jousseaume (1889; Mem. Soe. Zool. France, II, p. 248, figs. 
ix-12, 13) and T'richia rojasi Jousseaume (l.c., p. 249, figs. 
ix-9, 10), both from Venezuela; although venezuelensis is 
described as hirsute, the figure of rojasi is the one that shows 


Occasional Papers of the Museum of Zoology 79 


the hairs. On the other hand, the shape of these two forms is 
closer to that of Thysanophora vanattar (see below). 

The sculpture of all of these species is somewhat similar to 
that in 7. fuscula (C. B. Adams), T. proxima Pilsbry, and 
T. canalis Pilsbry (cf. 1922; Oce. Papers Mus. Zool. Univ. 
Mich., no. 106, p. 56), but the subdiscoid shape of crinita and 
arubana is more like that of the generic type, T. conspurcatella 
(Morelet). The haphazard distribution of the long hairs is 
very peculiar, but these are only present in the young shells 
of arubana. 


Thysanophora vanattai, new species 


Type locality: (A4a) at base of northern escarpment, Seroe 
Canashito, Aruba. 

Distribution : Aruba; Baranca Alto (A2ce; very rare, 1 speci- 
men), Rooi Frances (A3; quite infrequent, 33 specimens), 
Seroe Canashito (A4; quite rare, 22 specimens). Under lime- 
stone rocks, in protected places. 

Shell (fig. xv-57): small, depressed turbinate, thin and 
translucent; very light horn-colored, almost white; epidermis 
thin, but usually incrusted heavily with reddish material and 
even small particles from the surrounding rocks. Whorls: 
43/,, quite evenly rounded; very gradually increasing ; suture 
well impressed; last whorl very slightly descending. Sculp- 
ture of last whorl: rounded growth-wrinkles, separated by fine, 
impressed lines ; crossed by more oblique striae, which are very 
weak and short. Embryonic whorls: 114; practically smooth, 
although with extremely minute, irregular punctations, 
which have a slight tendency towards spiral arrangement. 
Umbilicus: open, circular; about 14 the major diameter of the 
shell. Aperture: slightly oblique, broadly elliptical. Peri- 
stome : incomplete, simple and sharp. 

Radula (fig. xiv-50): C/3, L6/2, M2/3 + (4/4-++) + 1/1; 
or 10-6-1-6-10. Central: symmetrical, with rather slender, 
sharp cusps. First lateral: similar but asymmetrical, with 
entocone lacking; on the inner side of the mesocone is a narrow 


80 University of Michigan 


expansion, which becomes broader in the successive laterals 
until, in the 7th tooth, it develops a small cusp. This cusp be- 
comes larger on the 8th, while, in the 9th, the ectocone also 
becomes bifid. In the outer marginals, the ectocone breaks up 
still more, but the inner two cusps remain about the same. 
The outermost tooth is a mere denticle. Jaw (fig. xiv—51): 
thin, slightly arcuate; consists of 25 to 27, transverse plates, 
which serrate the margins, but appear to be well fused 
together. 


Measurements 
altitude maj. diam. min.diam. lt.apert. diam.apert. whorls 
2.58 165 (4.19) 150 (3.92) 55 (1.43) 135 (1.95) 434 


This species appears to be somewhat similar to the small 
Thysanophorae of Porto Rico and the northern Lesser Antilles, 
such as 7. subaquila (Shuttleworth). It is smaller and has a 
relatively larger umbilicus. 


BULIMULIDAE 


Drymaeus virgulatus (Férussac) 


Heliz elongata ‘‘Bolten’’ Roeding (1798; Mus. Bolt., 107). Helia 
virgulata Férussac (1821; Tabl. Syst., 54; Hist. fig. exlii, 1-7). 
Bulimulus apiculatus Gray (1834; P. Z. S., 66); the form with the 
bluish apex. H. ludovica ‘‘Rang’’ Beck (1837; Index, 66); reddish, 
unicolor form. Bulimus elongatus Bland (1861; Ann. Lye. Nat. Hist. 
N. Y., VII, 143) ; first Bonaire record. Bulimus elongatus Bland (1866; 
Amer. J. Conch., II, 143); first Curagao record. Bulimulus elongatus 
Smith (1898; Proc. Mal. Soc., ITI, 114); first Aruba record; collected 
by Hartert. 


Type locality: Porto Rico. 

Distribution: Porto Rico; northern Lesser Antilles; Vene- 
zuela? Curacao: in the richer places throughout the island 
(C1, 2, 4, 9, 1led, 12a, 18-17, 20; Cb16, 17). Aruba: Seroe 
Canashito and Rooi Taki (A4a, 2c). Bonaire: in the better 
wooded places throughout the island (B3, 4, 5, 6, 7, 8; Bb3, 7). 
Usually found cemented to the trunks or branches of the larger 
trees; often found on the gumbo limbo (Bursera gummifera) 


Occasional Papers of the Museum of Zoology 81 


and guayacan. Quite independent of the nature of the under- 
lying rocks. 

Férussae apparently based his typical form on Lister’s fig- 
ure 2. This shows a rather small ‘specimen with the color- 
varices broken by a tendency towards spiral bands. The fig- 
ure (a) of Chemnitz (vol. IX, pl. 134, fig. 1225) shows a 
specimen with only the varices, while his figure (b) shows 
both these and the spiral bands. 

In the series from the Dutch Leeward Islands, the color 
variation may be analyzed as follows: 

Apex: every intergradation between pure white and quite 
dark, lilac-blue (apiculatus). Although both extremes com- 
monly occur in the same locality, the colonies usually tend 
towards a preponderance of one or the other of these color 
forms. For instance, 97 per cent. of the specimens from 
Bonaire have dark apices, while none of those from Aruba 
show this tendency. Both forms are present in about equal 
numbers in Curacao as a whole (53 per cent. white, 47 per 
cent. blue). As a rule, the blue-tipped shells are more strik- 
ingly banded, although some of the shells of this form from 
Bonaire are otherwise colorless. 

Ground color: dead white in all of my specimens, but in 
some the band: coloration is so diffuse as to tinge the last whorl 
with light chocolate. Shells in one lot in the A. N. S. P. (no. 
2430, collected by Raven), from Bonaire, are salmon-pink and 
almost unicolor (ludovica). 

Bands: in this species the spiral and growth sets of color 
banding seem to struggle for dominancy. When present, 
either set may be diffuse or sharply marked, narrow or broad, 
and may vary in color from light reddish brown to deep choco- 
late with a purplish tinge. Ten per cent. of my specimens are 
without bands. Nine per cent. have numerous, quite regular 
varices, which are usually quite narrow and sharply marked. 
This type of coloration is especially conspicuous near Land- 
huis Hato, Curacao (C1led). All of the shells show a distinct 
tendency to increase the density of the pigmentation during 
the periods of slower growth (this, by the way, is a character- 


82 University of Michigan 


istic of most of the species in the Dutch Leeward Islands), 
and, as a result, the spiral bands are seldom continuous for 
any distance. Forty per cent. of the shells may be classed as 
flammulated. In this group are included the shells in which 
the varices are broken by non-pigmented spiral zones, and also 
those in which the continuous or broken varices are flammu- 
lated by spirally arranged blotches of color. In 30 per cent. 
of the shells, the broken, spiral bands and varices are about 
equally prominent, so as to form grid markings. Twelve per 
cent. may be classed as spirally banded, although only one 
specimen (from B8) has truly continuous, spiral bands for 
any distance. Usually, the spiral bands are lightest where the 
growth lines are farthest apart; this produces a series of con- 
tiguous, oblong blotches, each of which shades from light to 
dark in the direction of growth. 

Peristomal callus and edge of columella: usually white, 
bordered internally by a dark, chocolate band; sometimes the 
entire callus is colored practically as dark as the interior of 
the aperture. 

Interior of the aperture: almost white to dark chocolate, 
usually darker than the exterior of the shell, but varying in 
intensity of color with it. Commonly the spiral bands and 
varices are also darker and broader than on the exterior. 

In addition to the color variation, the shells vary consider- 
ably in size and shape, and in solidity. The shells from Aruba 
are the lightest and smallest, while those from Bonaire are the 
largest and most solid. The comparatively small numbers of 
specimens make the following dimensions studies less trust- 
worthy than in some of the other species. In addition, on ac- 
count of the large numbers of immature specimens that. were 
present among the adults, only the larger specimens were col- 
lected, and the sets are scarcely random lots. 

The remarkably discontinuous distribution of D. virgulatus 
gives rise to the suspicion that it has been introduced into 
Curacao by commerce; it is noteworthy that this species also 
occurs in St. Martin and St. Eustatius, two of the islands of 
the northern Lesser Antilles, which are also part of the Nether- 
lands Colony of Curacao (cf. Vernhout, 1. c., p. 184). 


Occasional Papers of the Museum of Zoology 


83 


Table VIII. Dimensions of Drymaeus virgulatus 
Comparison of Stations 


place nos. 

Lister; fig. 2 
Férussac ; 

pl. 142B, 1 

Cl 26 
C2 25 
C4 30 
C9 + 
Clled 21 
Cl2a 13 
C13 7 
Total; 


s. Curacao 126 


C14 3 
C15, 17 24 
C16 11 
Cb16, 17 2 
C20 8 
Total; 

n. Curacao 48 
A2e 5 
A4a 5 
Total; 

Aruba 10 
B3 to 7; 

Bb3, 7 64 
Bs 7 
Total; 

Bonaire 71 


Cl. Quite infrequent. 
Quite infrequent. 
C4. Frequent, on trees and even the crotons. 


C2. 


sharply marked. 


whorls 


6+ 


7 
6.5 (6.0-7.0) 
6.4 (6.0-7.0) 
6.9 (6.5-7.5) 
6.8 (6.5-7.0) 
6.5 (6.0-7.0) 
6.7 (6.5-7.0) 
6.6 (6.0-7.0) 


6.6(6.0-7.5) 
6.3(6.3-6.5) 
6.6(6.3-7.3) 
6.6(6.3-7.0) 
6.8(6.5-7.0) 
6.8 (6.5-7.0) 


6.6 (6.3-7.3) 
7.1(7.0-7.3) 
6.9 (6.8-7.0) 
7.0 (6.8-7.3) 


7.0(6.5-7.8) 
6.6(6.3-7.0) 


6.9 (6.3-7.8) 


altitude 
25.6 


32.1 
25.5 (23.1-29.3) 
26.0 (23.0-29.9) 
25.9 (22.5-29.8) 
26.6 (26.2-26.8) 
25.9 (23.6-28.3) 
26.3 (23.4-27.9) 
26.6 (24.2-29.6) 


26.1 (22.5-29.9) 
25.5 (24.0-27.0) 
26.6 (23.8-29.2) 
27.8 (26.6-30.6) 
28.4 (27.5-29.3) 
27.4(25.4-29.1) 


27.1(23.8-30.6) 
25.6 (24.3-27.4) 
23.8 (22.7-25.3) 
24.7 (22.7-27.4) 


28.6 (24.5-32.2) 
27.2 (25.2-28.2) 


28.4 (24.5-32.2) 


C9. Very infrequent. Sharply marked. 


Clled. Infrequent. 


Cl12a. 


index 
41 


42 
46 (44-50) 
47 (44-50) 
45 (42-49) 
46 (45-48) 
47 (44-50) 
46 (45-48) 
47 (43-49) 


46 (42-50) 
48 (46-50) 
46 (43-49) 
47 (45-50) 
46 (44-48) 
45 (44-46) 


46 (43-50) 
46 (42-47) 
47 (46-48) 
46 (42-48) 


45 (40-50) 
47 (45-48) 


45 (40-50) 


Bright, and sharply marked. 
Dull and diffuse coloration. 
Sparsely but quite 


Sharply and finely marked. 
Quite infrequent, on larger trees at base of the escarpment. 


maj. diam. 
10.6 


13.6 
12.1(11.1-12.8) 
12.2(11.4-13.4) 
11.8(10.9-13.1) 
12.2(11.9-12.8) 
12.2(10.6-13.0) 
12.0(10.7-12.7) 
12.2 (11.5-12.9) 


12.1(10.6-13.4) 
12.2(11.9-12.5) 
12.1(11.0-13.3) 
13.1(12.5-13.8) 
13.0(12.9-13.2) 
12.2 (11.4-13.1) 


12.5 (11.0-13.8) 
11.6(11.3-12.1) 
11.2(11.0-11.7) 
11.4(11.0-12.1) 


12.9(11.0-13.8) 
12.6(12.1-13.6) 


12.9(11.0-13.8) 


Peristomal callus heavy; shells rather solid. Dark, broad sharply-marked 
bands and varices. 


C13. Infrequent, mainly at base of northern escarpment. 


diffuse coloration. 


C14. 


Very infrequent. 


Similar to C13. 


Bleached, 


84 University of Michigan 


C15 and Cl7a, lots mixed. Frequent. Shells rather solid, with heavy 
peristomal callus. Coloration diffuse, to dark and sharply marked. 

C16. Frequent. Shells solid with heavy callus. One specimen has the 
heaviest callus in the sets; the shell is 2 mm. thick on the palatal wall. 
Dark bands and varices. 

Cb16, 17. Rare to infrequent; occurs in the richer valleys throughout 
the region of older rocks from Campo Sint Kruis to the Tafelberg of Sint 
Hyronimus. Quite heavy, with diffuse markings. 

C20. Infrequent. Similar to Cb17. 

A2e. Very rare; only found on Cereus in Rooi Taki. Thin and dull- 
colored. 

A4a. Rare; on trees at base of northern escarpment. Similar to A2e. 

B3-7; Bb3, 7. Rare to frequent. In the richer localities, both over 
limestone and in the interior region. The largest shells collected; solid 
with heavy peristomal callus. Usually rather dull colored. 

B8. Quite infrequent. Somewhat smaller than the preceding. 


From the comparison of dimensions (Table VIII), it will be 
seen that typical D. virgulatus, as judged by the figures, is a 
considerably more slender shell than that represented by the 
mean of the Curacao specimens, and, in fact, corresponds to 
the most slender shells from the Dutch Leeward Islands. It 
would be interesting to compare a large lot from Porto Rico. 

The radular formula (fig. xiv-52) of a specimen from Seroe 
Papaja (Cl1l1c) is: 86-1-86. The central is asymmetrical and 
tricuspid. Almost all of the laterals are also tricuspid, but, 
as is usual in the genus, the ectocone is very variable and may 
be entire and bifid in two consecutive teeth; as a rule, the 
outer teeth are more variable than the inner, but the first 
lateral itself may be 4-cusped, while the 83rd tooth is com- 
monly tricuspid. The transverse rows run obliquely back- 
wards to the 54th tooth, which is peculiarly elongate and lacks 
the entocone ; with the 55th, the rows curve abruptly forward so 
that the entire row has the shape of a broad W (see line under 
scale in figure). In the 1st to the 53rd teeth, the entocone is 
markedly larger than the ectocone and is quite widely separated 
from the mesocone, but the teeth beyond the 54th are more 
nearly symmetrical. In addition, the bases, in each portion, 
are almost parallel to the direction of the row, so the teeth of 
the outer and inner limbs are quite markedly different in ap- 


Occasional Papers of the Museum of Zoology 85 


pearance. The outermost teeth are strongly compressed trans- 
versely. This radula is very similar to that figured by Pilsbry 
(1902; Man. Conch., XIV, fig. LX-16) for D. interpunctus 
(Martens). 


Oxystyla maracaibensis imitator Pilsbry 
(1899; Man. Conch., XII, p. 140, fig. xxx, 49-54) 


Type locality: near Maracaibo, Venezuela. 

Distribution: Maracaibo, Venezuela, and Santa Marta, Co- 
lombia, to Peru. Aruba: subfossil in the aloe fields on the 
western side of the island. Four specimens collected about 1 
km. south of Seroe Canashito. From the localities listed, this 
appears to be a desert form. 

Although all of my specimens are badly bleached, from one 
to three spiral bands and two or three varices can be made out. 
One specimen is almost a perfect duplicate of fig. 54 in 
Pilsbry (1. c.). 


Measurements 
altitude maj. diam. min.diam. alt.apert. diam.apert. whorls 
45.4 61 (27.5) 52 (23.7) 56 (25.5) 69 (17.6) 7 
ACHATINIDAE 


Genus Neosubulina, subgenus s. s. 


Neosubulina Smith (1898; P. Mal. S., III, p. 115); monotype Neo- 
subulina harterti Smith, from Bonaire. _ 


Shell: ovate-turrite ; light-colored and translucent. Whorls: 
numerous. Growth-wrinkles of last whorl: well defined. 
Embryonic shell (dissected out from parent): 214 to 2%4 
whorls; with delicate and regular growth-wrinkles crossed by 
spiral striations, which extend to the very apex. Umbilicus: 
rendered imperforate by expansion of columellar-parietal 
callus. Peristome: thin, sharp, incomplete; lower palatal wall 
broadly and slightly emarginate; with a twisted thickening 
around the columella, which is present even in the embryonic 
shell; parietal wall with a spiral lamella in the adult. Repro- 


86 University of Michigan 


duction: ovoviviparous; eggs relatively very large, with a 
white, granulate capsule. Radula: very similar to that of 
Opeas, but with multicuspid marginals. 

In this genus (see N. harterti), the inner laterals are tri- 
cuspid and almost symmetrical, while the outer ones are re- 
duced in size and slightly more tilted inwards; in Leptinaria 
(cf. Pilsbry ; 1907; Man. Conch., XVII, fig. xli4), the ento- 
cone is very much reduced on the inner laterals, while the 
outer ones are extremely elongate. Although the lingual 
armature is known only in the typical group, I believe that 
Neosubulina includes three subgenera: Pelatrinia Pilsbry (l. 
c., p. 324), monotype Leptinaria helenae Pilsbry from Vene- 
zuela; Neosubulina s.s. (redeseribed above) from the Dutch 
Leeward Islands; and Ischnocion Pilsbry (Il. c.), monotype 
Leptinaria triptyx Pilsbry from Colombia. In the first of 
these, the shell is subacuminate-turrite; the embryonic whorls 
are vertically striate; and only the columellar twist is present 
in the adult. In the second, the shell is ovate-turrite; the 
embryonic shell is vertically and spirally striate; and a spiral 
lamella is present on the parietal wall of the adult. In the 
third, the shell is subeylindric-turrite; the embryonic whorls 
are practically smooth; and the adult develops a palatal fold 
in addition to the columellar twist and the parietal lamella. 


Neosubulina harterti Smith 
(1898; Proce. Mal. S., III, p. 115, fig. IT) ; collected by Hartert 


Type locality: Bonaire; probably near Kralendijk (B1, 2). 

Distribution: Bonaire; in the richer localities (B3-6), 
buried deeply in limestone talus. The species of this genus 
inhabit the rotten mould that fills the deeper crevices of the 
detritus ; they seldom occur among the cleaner rock fragments 
near the surface, although the latter is the stratum where most 
of the other genera were found. For this reason, it is very 
difficult to estimate the abundance. 

Shell (fig. xvi-61) ; light horn-colored, quite transparent ; 
tapers quite regularly from last whorl to apex. Whorls: 9 
(maximum observed); later whorls elongate, slightly and 


Occasional Papers of the Museum of Zoology 87 


evenly convex; suture oblique, shallow except near the apex. 
Seulpture of last whorl: growth-wrinkles fine, regular and 
thread-like, vaguely and very lightly crenulated by a few, 
spiral striations. Aperture: elongate-ovate, with long axis 
slightly oblique to that of shell. Peristome: columellar trunca- 
tion slight in adults but relatively prominent in the embryonic 
shell; parietal lamella compressed, present as a thin lamella 
on the central axis of the last but dying out on the penultimate 
whorl of the adult, present as a fine, internal thread in a young 
shell of 514 whorls, not developed in embryonic shell (fig. 
xv1—58). 

Radula (fig. xiv-53): C/3, L7/3, M4/-+ (10/4-++): or 
21-1-21. Very similar to that of Opeas beckianum (cf. 1923; 
this series, no. 135, fig. I-6), but the cusps of the inner laterals 
are broader and heavier; the teeth in each transverse row de- 
crease more rapidly in size towards the outside; and, in the 
marginals, the entocone (first), ectocone and mesocone become 
subdivided into minor cusps (all usually tricuspid in Opeas). 

Jaw (fig. xvi-64): thin, transparent, arcuate, crossed by 
numerous, well-impressed lines, which separate rounded, gran- 
ulate riblets. 


Measurements 
alt. maj.diam. alt.apert. diam.apert. whorls 
embryo (fig. 58) 1.8  60(1.07) 58 (1.04) 53 (0.55) 244 
largest, B6 (fig. 61) 11.5 20(2.33) 22 (2.57) 54(1.39) 9 
Smith (1898) 9.0 25(2.25) 22(2.0 ) 8 


~As Smith (l.c.) was apparently unfamiliar with the de- 
scription of N. gloynii, he did not differentiate the two species. 
As a result, his species would be unrecognizable without the 
locality. The absence of the parietal lamella in the embryonic 
shell indicates that NV. harterti is more distantly related to N. 
gloynii and N. scopulorum than they are to each other. The 
specimens from the northern localities on Bonaire (my meas- 
urements and figures) average slightly larger than do those 
from the more southern stations (probably typical hartertt). 
Although considerable search was made for it, this species was 
not obtained on Klein-Bonaire. 


88 Unversity of Michigan 


Neosubulina gloynu (Gibbons) 


Cionella gloymi Gibbons (1879; J. of Conch., II, p. 135, fig. I-1); 
W. G. Binney (1883; Ann. N. Y. Acad. Sci., III, p. 101, fig. vii-3) ; 
radula. Leptinaria gloyni minuscula Pilsbry (1907; Man. Conch., 
XVIII, p. 323, fig. xlvii-18); small form. Leptinaria gloynu H. B. 
Baker (1923; this series, no. 137, p. 5, figs. I, 1-3). 


Type locality: ‘‘St. Ann’s, numerous under stones,”’ 
Curacao; probably the Schaarlo (C5c), back of Willemstad. 

Distribution : Curacao; in the richer localities, both on lime- 
stone (C1-—6, 11, 12, 15, 17, 20) and in the central region (Cb2, 
6, 17, 20) ; not found near the shore but reaching an altitude 
of 1,200 feet on Sint Christoffelberg. Habits as in hartertt. 
This is the only species that was found in the groves of poison- 
ous manzalienja trees along the inland water-courses; it was 
found in only one such locality (Cb2), although the thick 
layers of fallen leaves in these places look like rather favorable 
localities for land molluses. 3 

Shell (fig. xvi-62): cloudy pearl-colored, usually more 
opaque than the other two species; averages larger and tapers 
more abruptly near the apex than does harterti. Whorls: 934 
(maximum observed) ; later whorls slightly stouter (especially 
near the middle of the shell) and lower than in harterti, each 
convex above the middle and tapering and flat-sided below so 
that they appear to telescope into each other; suture more pro- 
nounced and slightly less oblique. Sculpture of last whorl: 
growth-wrinkles coarser but more obscure and irregular than 
in harterti, also more oblique and markedly arcuate just below 
the suture; spiral sculpture quite absent. Aperture: slightly 
more elongate and with long axis more oblique to that of shell. 
Peristome: columellar truncation of the adult more pro- 
nounced ; parietal lamella heavier than in harterti, developed 
as a marked angulation (cf. 1923, 1. c., fig. 3) on the central 
axis of last and penultimate whorls, low but distinct on the 
smaller embryonic shell (fig. xvi-59) ; lower palatal emargina- 
tion more pronounced than in hartertt. 

In the more barren localities, the shells are usually somewhat 
smaller and may be included in the form minuscula (Pilsbry). 


Occasional Papers of the Museum of Zoology 89 


Otherwise the species appears quite constant throughout the 
island. 


Measurements 
alt. maj.diam. alt.apert. diam.apert. whorls 
embryo (fig. 59) 1.6 62(0.97) 58 (0.91) 59 (0.53) 21% 
largest, C11 (fig.62) 12.7 21(2.65) 22 (2.84) 52 (1.48) 934 
Gibbons (1879) 13 23 (3) 9-10 


Neosubulina scopulorum, new species 


Type locality: (A8) at base of right wall of entrance to 
Rooi Frances, Aruba. 5 

Distribution: Aruba; richer localities on the limestone 
(A2-4). Habits as in harterti. 

Shell (fig. xvi-63) : bright horn-colored, quite transparent ; 
tapers abruptly near the apex as in gloynii, but is also con- 
stricted noticeably just below the middle; attains a much 
larger size (and greater number of whorls) than do the other 
two species. Whorls: 1134 (maximum observed) ; apical and 
subapical quite similar to gloynii; those just below the middle 
markedly elongate, convex below the suture but flat-sided and 
tapering below the convexity so that the telescopic appearance 
is even more marked than in gloyniz; suture more pronounced 
and oblique. Sculpture of last whorl: growth-wrinkles more 
closely-spaced and angular than in gloynu, but coarser than in 
harterti, slightly less oblique than in the former but similarly 
arcuate just below the suture; spiral sculpture only indicated 
by broad and obscure constrictions. Aperture: shaped much 
as in gloyni, but with long axis slightly less oblique to that. of 
shell. Peristome: columellar truncation and parietal lamella 
of adult higher and larger than in gloynit, relatively as pro- 
nounced in the larger embryonic shell (fig. xvi-60) as in the 
adult; lower palatal emargination as in gloyndi. 

Radula: C/3, L9/3, M12/4 + ; or 21-1-21. Very similar to 
that of harterti, but the laterals decrease in size more gradu- 
ally and the first 9 teeth are tricuspid. Jaw: very similar to 
hartertt. 

The embryonic shells figured (figs. xvi, 58-60) were all dis- 
sected out of the bodies of the parent shells. As large eggs 


90 University of Michigan 


were rarely found in specimens from Curacao (June), but 
were more common in those from Aruba (July), and were 
present in a large proportion of the individuals from Bonaire 
(August), it seems probable that the eggs are laid near the 
beginning of the wet season. The adult figured (figs. xvi-61, 
62, 63) is, in each species, the largest specimen collected. In 
all of the species, the spire is often curved out of line with the 
lower whorls, but this asymmetry is most common in N. 
scopulorum. 
Measurements 

alt. maj.diam. alt.apert. diam.apert. whorls 
Embryo (fig. 60) 1.7 62(1.07) 52(0.90) 60 (0.54) 2% 
Type, largest (fig.63) 16.9 16(2.69)  18(2.99)  52(1.57) 11% 


UROCOPTIDAE 


Brachypodella raven ravem (Crosse) 


Cylindrella raveni ‘‘Bland’’ Crosse (1872; J. de C., XX, 157) ; Crosse 
and Bland (1873; J. de C., X XI, fig. I-4) ; collected by Raven. Brachy- 
podella raveni Pilsbry (1903; Man. Conch., XVI, fig. ix—14); radula. 


Type locality: Curacao; probably the Schaarlo (C5e), back 
of Willemstad. 

Distribution of species: Curacao, Aruba; under limestone 
rocks near the surface, in all but the most barren localities. 

Distribution of subspecies: Curacao; from New Port north 
to Landhuis Hato (at least C1, 3, 4, 5, 6, 7, 11, 12). 

Shell (fig. xvii-67): brownish in color with lighter ribs, 
opaque; subacuminate-turrite, greatest diameter at 214 to 3° 
whorls from aperture. Whorls: 14 (in a rather small speci- 
men), of which about 9 are usually retained; moderately con- 
vex with well-impressed sutures; last whorl with basal carina 
below a peripheral emargination. Sculpture of later whorls: 
fine, but well-marked growth-riblets, which are usually as wide 
or wider than their interspaces and slightly crested over the 
basal carina. Embryonic whorls (fig. xvii-73) : apparently 3, 
of which the 2nd is as broad or broader than the 3rd; thin, 
translucent, amber-colored; sculpture of fine, closely-spaced, 


Occasional Papers of the Museum of Zoology 91 


regular growth-riblets. Plane of aperture considerably in 
front of periphery of last whorl. , 

Radula (fig. xvi-65) : C/3-4, L2/2, M7/2 + 1/0; or 8-2-1- 
2-8. Central: with very slender base; the outer cusps are 
large, but one or two, very variable, smaller cusps lie between 
them; this seems to indicate a tooth that is becoming bicuspid 
by the suppression of the central cusp of the tricuspid type. 
The accessory cusp of the inner lateral is aculeate in shape, 
while that of the outer lateral is broader and more hoe-shaped. 
The slender bases of the small marginals are very irregular at 
their posterior ends, while the lateral corner of the anterior 
margin is produced into a sharp point. 124 rows of teeth 
counted in a radula which lacks the unformed, posterior 
portion. 

Jaw: consists of about 44, thin, slender, subrectangular, 
overlapping plates, which are shorter near the center where 
they form a triangle. , 


Measurements 
Shell Spire Aperture Whorls 
alt. maj.diam. maj. diam. alt. diam. 


C11, entire 
(fig. 67) 8.98 29(2.63)  24(2.17) 16(1.46) 104(1.52) 14 
C5, largest 7.83 34(2.64)  28(2.76) 21(1.68) 104(1.74) 8 


Typical ravemi attains the largest size and has the most 
closely spaced riblets of any of the forms in the Dutch Lee- 
ward Islands; all of these are placed by their apical sculpture 
in the mainland group of Brachypodella s.s. (cf. Pilsbry, 
op. cit.). As will be noted from the measurements of this and 
the following forms, the few complete shells obtained are con- 
siderably below the maximum size. The largest shells also 
appear to have developed the greatest number of whorls. 


Brachypodella raveni sanctaebarbarae, new subspecies 


Type locality: (C2a) at base of northern escarpment of the 
Tafelberg of Santa Barbara, southern Curacao. 

Distribution: Curacao; only found at the base of the north- 
ern and western escarpments of this Tafelberg (C2). 


92 University of Michigan 


Shell (fig. xvii-68) : considerably smaller and much thinner 
and more polished and translucent than raveni. Growth- 
riblets of later whorls much lower, more rounded, and more 
widely spaced. Aperture relatively larger. Otherwise as in 
ravem. 


Measurements 
Shell Spire Aperture Whorls 
alt. maj. diam. maj. diam. alt. diam. 


C2, type 
(fig.68) 7.75 30(2.32)  26(2.00) 18(1.38) 108(1.49) 13% 
C2, largest 7.76 28(2.84)  28(2.19) 23(1.77) 103(1.83) 8 


The several hundred specimens collected from the type 
locality are all quite distinct from typical ravem, but they do 
intergrade slightly with those from near New Port (C1), 
which are connected by intermediates with those from the more 
northern localities. With equal reason, these two subspecies 
could be considered as separate species which produce hybrids 
at their point of contact. 


Brachypodella raveni knipensis, new subspecies 


Type locality : (C17a) Seroe Djerimi, northern Curagao. 

Distribution: Curacao; north of Seroe di Boca (at least 
Cp, 11, 20). 

Shell (fig. xvii-69): averages considerably smaller than 
ravem; greatest diameter at about the 3rd whorl above the 
aperture. Carina and peripheral emargination carried higher 
on the whorls. Growth-riblets of later whorls more widely 
spaced and slightly heavier. Otherwise as in ravent. 


Measurements 
Shell Spire Aperture Whorls 
alt. maj. diam. maj. diam. alt. diam. 


C17, type 
(fig. 69) 7.97 30(2.39)  26(2.05) 16(1.27) 118(1.50) 138% 
C17, largest 7.24 36(2.61)  29(2.10) 21(1.51) 107(1.61) 8 


Occasional Papers of the Museum of Zoology 93 


Aperture formation in Brachypodella appears to be 
hastened’ by injury and specimens with peculiarly distorted 
peristomes are not uncommon. In the present subspecies, 
quite 2 number of the specimens, although apparently unin- 
jured, are almost perfectly conical, with the last whorl well 
rounded and larger than any of the preceding ones; a single 
specimen (fig. xvii-72) also has the peristome adnate to the 
last whorl. 


Brachypodella raveni arubana, new subspecies, 
and form sinistrorsa, new 


Type locality: (A2b) Seroe Pretoe, between Roois Hundoe 
and Spoki, Aruba. 

Distribution: Aruba, under limestone rocks. All of the 
specimens from A2, and A3, and 93 per cent. of the 105 shells 
from A4 are dextral (typical form) ; 7 per cent. of those from 
Seroe Canashito (A4), and all from the shore-cliffs between 
Perkietenboseh and Kralendijk (A5, quite rare) are sinistral 
(form sinistrorsa). 

Shell (xvii-70) : similar in size to ravent, but usually more 
cylindric; greatest diameter usually above the 3rd whorl; sur- 
face more highly polished. Whorls: 1414 (in a medium-sized 
specimen), of which about 10 are usually retained; later 
whorls less convex than in typical raveni; carina and periph- 
eral emargination carried much higher on the shell. Growth- 
riblets of last whorl: usually closely-spaced, but much lower 
and more rounded than in ravemi; carinal crests poorly de- 
veloped or absent. Peristome: reflection broader than in 
raven. 

Radula and jaw: similar to raveni, but the intermediate 
cusps of the central tooth are usually smaller. 


Measurements 
Shell Spire Aperture Whorls 
alt. maj.diam. maj. diam. alt. diam. 


A4, complete 8.85 24(2.39)  22(1.99) 18(1.59) 107(1.70) 14% 
A2, type, larg- 
est (fig.70) 8.32 32(2.67)  26(2.19) 21(1.75) 101(1.76) 8 


94 University of Michigan 


The specimens from southern Aruba are quite distinct from 
any of the other subspecies, but those from the central por- 
tions are usually smaller, often have more distinct riblets, and 
intergrade slightly with knipensis. The sinistral shells are 
similar in sculpture and size to the dextral specimens from 
Seroe Canashito (A4), but their definite distribution seems 
to require recognition as at least an incipient local race. The 
type locality of form sinistrorsa is the top of the low shore- 
cliffs just south of Kralendijk (A5c). 


Brachypodella gibbonsi, new species 

Cylindrella raveni Gibbons (1879; J. of C., I, 340); first Bonaire 
record. 

Type locality: (B3) base of western escarpment of Mon- 
tagne, Bonaire. 

Distribution: Bonaire (B3—6, 8), Klein-Bonaire (K1) ; un- 
der limestone rocks. 

Shell (fig. xvii-71) : considerably smaller, more acuminate, 
and lighter in color than raveni; greatest diameter at the 
penultimate whorl. Whorls: 1234 (in a medium-sized speci- 
men), of which about 8 are usually retained; more convex and 
with deeper sutures than raveni; last whorl relatively more 
elongate. Growth-riblets of later whorls: much higher, heay- 
ier, and more distant than in raveni; expanded into heavy, 
white crests over the basal carina. Plane of aperture about 
on a level with the periphery of the last whorl (7. e., although 
the last whorl is equally tangential, it is not carried as far for- 
ward as in raveni). Peristome: heavier. Embryonic whorls 
and other characters as in ravent. 

Radula (fig. xvi-66) and jaw: very similar to ravent, but 
the centrals are usually bicuspid although a rounded projec- 
tion is sometimes developed in the shallower notch. 148 trans- 
verse rows counted in a complete specimen. 

This species has more distant riblets than B. raveni sanctae- 
barbarae, while they are heavier than those of the subspecies 
knipensis. Although this species does not intergrade with 
either, it appears more closely related to the northern and 
southern forms than to typical raveni of central Curacao. 


Occasional Papers of the Museum of Zoology 95 


Measurements 
Shell Spire Aperture Whorls 
alt. maj.diam. maj. diam. alt. diam. 


B3, type, en- 
tire(fig.71) 7.64 34(2.61)  26(2.01)  20(1.50) 99(1.49) 12% 
B3, largest 7.10 37(2.61)  31(2.22) 22(1.56) 100(1.56) 8 


Microceramus bonairensis bonairensis (Smith) 

Pineria bonairensis Smith (1898; Proc. Mal. Soc., III, p. 114, fig. I) ; 
collected by Hartert. 

Type locality: Bonaire, probably near Kralendijk (B1, 2). 

Distribution of species: Bonaire, Klein-Bonaire, Curagao, 
and Aruba. Under superficial limestone slabs; this species is 
apparently next to the Pupillidae, among the smaller shells, 
in its ability to withstand aridity. 

Distribution of subspecies: Bonaire (B1-8) and Klein- 
Bonaire (K1). 

Shell (fig. xvii-74): ovate-lanceolate; penultimate whorl 
slightly the broadest. Color: light corneous, marked with 
opaque, milky-white, irregular patches. Whorls: about 9; 
quite convex; suture well marked. Sculpture of later whorls: 
regular, almost contiguous, quite prominent, rounded growth- 
riblets, which are not much obscured by the relatively incon- 
spicuous, opaque thickenings; basal whorl with slight, spiral 
angulation, which is first evident just below the parietal angle 
of the aperture. Embryonic whorls: corneous; convex, with 
impressed sutures; first half-whorl smooth and polished, re- 
mainder with minute, regular, well-marked, and quite closely- 
spaced growth-riblets. Umbilicus: distinctly rimate. <Aper- 
ture: subcirecular, but distinctly broader than high. Peris- 
tome: whitish; slightly thickened and reflected; incomplete ; 
thickened and most reflected on the columellar wall. Colu- 
mella inside of whorls: slender, with a slight, spiral thickening. 


Measurements 
Shell Aperture Whorls 
alt. maj. diam. alt. diam. 
Type (Smith, 1898) 6.0 42(2.5 ) 25(1.5 ) 8% 


Fig. 74 (B2) 6.49 49(3.17) 27(1.74) 112(1.95) 8% 


96 University of Michigan 


Typical bonairensis (Table IX, B2) from southern and cen- 
tral Bonaire (B1, 2, 7) and Klein-Bonaire (K1) is quite dis- 
tinet in form and sculpture from any of the other subspecies. 
However, those from the more northern localities (B5, 6, 8) 
are more slender and elongate (Table IX, B8), and the growth 
riblets of the lower whorls are weaker and less regular, so 
these shells intergrade somewhat with the series from Curacao, 
especially with the lots trom the more northern localities of 
that island. 

In Table IX, the minor diameter is used because of its 
greater constancy and on account of the difficulty in the meas- 
urement of the major diameters of these small shells by means 
of calipers. Throughout this paper, all measurements that 
involve two decimal places were made from camera-lucida 
drawings under considerable magnification. It will be noted 
that I did not obtain any specimens as small as Smith’s meas- 
urements would indicate, but this may be due to the fact that 
his figures are only to the nearest half-millimeter. 


Table IX. Dimensions of Microceramus bonairensis 
Place Nos. Whorls Altitude Index Min. Diam. 
B2 20 9.0( 8%4- 93%) 7.0(6.3— 7.9) 42(37-44) 2.9(2.8-3.1) 
B8 18 9.7( 834-101%4) 7.7(6.5— 8.7) 37(31-43) 2.8(2.7-3.0) 
C5 94 10.0( 834-1114) 8.0(6.4-10.2) 36(32-42) 2.8(2.6-3.3) 
C17 50 =9.5( 834-10%) 7.2(6.3— 9.2) 37(32-41) 2.7(2.5-3.1) 
A4 56 10.8(10 -12%4) 8.6(7.1-10.0) 33(30-36) 2.8(2.5-3.2) 


Microceramus bonairensis curacoanus H. Burrington Baker 


Macroceramus inermis Gibbons (1879; J. of C., II, 136) ; first Curagao 
record; W. G. Binney (1883; Ann. N. Y. Acad. Sci., ITI, 126); radula. 
Microceramus bonairensis cwracoana H. B. Baker (1923; Occ. Papers Mus. 
Zool. Univ. Mich.; no. 137, p. 6-7; figs. I-4, 5). 


Type locality: (C5e) Schaarlo, back of Willemstad, Curagao. 

Distribution : Curacao; under limestone rocks (C1-7, 11-13, 
17, 20). 

Shell (fig. xvii-75) : more elongate but slightly smaller than 
typical bonairensis. Color: light corneous to dark brown; 
opaque calluses more conspicuous than in preceding. Whorls: 
about 10; less convex and with shallower suture than in 


Occasional Papers of the Museum of Zoology 97 


bonairensis. Sculpture of later whorls: growth-riblets low, 
irregularly-spaced, often almost obsolete on last whorl; the 
calluses often obliterate the interspaces and may be decorated 
with faint, spiral thickenings; basal angulation slightly more 
prominent than in bonairensis. Aperture: subcireular. Other- 
wise as in bonmrensis. 


Measurements 
Shell Aperture Whorls 
alt. maj. diam. alt. diam. 
Type, fig. 75 (Cdc) 8.20 37(3.02) 22(1.84) 109(2.01) 10% 
Fig. 76 (C17) 7.20 37(2.66) 23(1.64) 108(1.76) 914 


The radula of this form has been described and figured in 
a former paper (1923; op. cit., fig. I-5). As indicated, it ap- 
pears closest to that of M. pontificus, but it differs from the 
radulae of the more northern species in its simply unicuspid 
central. The 105 transverse rows are almost straight. 

Typical cwracoanus (Table IX, C5) is distributed through- 
out the limestone portions of central and southern Curacao 
(C1-7, 11,12). In the more northern localities (C13, 17, 20), 
it is represented by a smaller form (Table [X, C17; fig. xvii— 
76) with slightly more regular growth sculpture. 


Microceramus bonairensis arubanus, new subspecies 


Type locality: (A4b) top of Seroe Canashito, Aruba. 

Distribution: Aruba; under limestone rocks (A2-5). 

Shell (fig. xvii-77) : more elongate than curacoanus. Color: 
lighter and more translucent, calluses less conspicuous than 
in preceding. Whorls: about 1034; more convex and with 
better impressed suture than either of the other subspecies. 
Sculpture of last whorl: growth-riblets more regular than in 
curacoanus but much lower and more rounded than in bonair- 
ensis ; calluses thin but widespread, usually with several spiral 
thickenings; basal angulation more noticeable than in the 
other subspecies. Otherwise as in curacoanus. 

This (Table IX, A4) is the largest form of the species, and 
also develops the greatest number of whorls. Its sculpture 


98 University of Michigan 


and form are much closer to curacoanus than that subspecies is 
to bonmrensis. 


Measurements 
Shell Aperture Whorls 
alt. maj. diam. alt. diam. 
Type, fig. 77 (A4) 9.96 33(3.32)  20(2.02) 104(2.10) 12% 


Table X. Type Measurements of New Forms of Cerion wa 
Shell Spire Aperture Whorls 
alt. maj.diam. diam. alt.’ diam. 

diablensis, 

fig. xvili—A2 (C12b) 19.4 40( 7.8) 40( 7.8) 30(5.7) 96(5.5) 11% 

hatoensis, 

fig. xvili-F'6 (Clld) 25.1 45(11.3) 45(11.2) 29(7.4) 99(7.3) 12% 

djerimensis, 

fig. xix-Al (Cl7c) 18.2 45( 8.1) 45( 8.1) 31(5.6) 95(5.3) 10%4 

knipensis, 

fig. xix-F5 (Cb17b) 25.2 44(11.1) 44(11.1) 32(8.1) 97(7.9) 11% 

arubanum, 

fig. xx-C3 (A2c) 21.2 47(10.0) 44( 9.3) 33(6.9) 91(6.3) 11% 

kralendijki, 

fig. xxi-A2 (B1) 18.9 46( 8.7) 44( 8.4) 32(6.0) 102(6.1) 10 

bonmrensis, 

fig. xxi-F'6 (B5) 26.9 42(11.3) 40(10.8) 32(8.7) 94(8.1) 12 


CERIONIDAE 


Cerion wva uva (Lin.) 


Turbo wa Lin. (1758; Syst. Nat., X, 765) ; description unrecognizable 
but the first reference to a figure (Pet. gaz. t., 27, f. 2) is this species. 
Cerion vulgare ‘‘Bolten’’ Roeding (1798; Mus. Bolt., 90); the reference 
to Knorr 6, t. 25, f. 4. Cerion apiarum ‘‘ Bolten’’ Roeding (I. ¢.) ; simply 
a reference to Turbo wa Gmelin. Pupa wa Schubert and Wagner 
(1829; Conch. Cab., XII, pl. 235, figs. 4122, 4123). Pupa wa Beck 
(1837; Index, 82); the first citation of Curacao as the habitat. Cerion 
woa Moerch (1852; Cat. Yoldi, 33). Turbo wa Hanley (1855; Ipsa 
Linn. C., 343); Linnaean specimens like figures in Conch. Cab. (see 
above). Cerion wa desculptum Pils. and Vanatta (1896; P. A. N.S. P., 
pp. 318, 328; fig. xi-1); form with reduced sculpture. 


Type locality : unknown; probably the Schaarlo (C5e), back 
of Willemstad, Curacao. 


Occasional Papers of the Museum of Zoology 99 


Distribution of species: Curacao; practically everywhere on 
the limestone, and also invading the region of the older rocks. 
Bonaire: as prevalent as on Curacao. Klein-Bonaire: every- 
where. Aruba: Perkietenboseh and Baranca Alto. Mainly on 
brush and eacti a short distance above the ground, but also 
on and under rocks. Very abundant in favorable localities; 
as many as 150 per square meter counted. 2,737 adults col- 
lected. 

Distribution of subspecies: Curacao; the southern portion, 
north of Kaap Sint Marie and Landhuis Hato (at least) ; 
mainly on limestone (C1—13), but also narrowly invades the 
region of the older rocks (Cb6, 10). 

Extreme variability appears to be a characteristic of this 
genus. In order to obtain some statistical idea of the amount 
and character of the variation in C. wva, the altitude, major 
diameter (exclusive of aperture), and the number of whorls 
were ascertained in all of the Curacao specimens collected. 
These data indicate that the variation may be immediately 
divided into two phases: the number of whorls and the dia- 
meter of the shell. The former would seem to be simply a 
function of the period of growth; while the latter expresses 
actual variation in size (dwarfing or gigantism). The dia- 
meter does not seem to be especially correlated with the num- 
ber of whorls; in other words, the larger shells do not appear 
to develop more whorls, although this is not strictly true of 
the extremes in size. Inside of each subspecies, on the other 
hand, the altitude is very closely correlated with the number 
of whorls (see Table XI). 

Table XII shows immediately that neither of these varia- 
tions is geographical; colonies with low (or high) means occur 
in widely separated places. The explanation is, I believe, 
purely ecological. The size of the shells (7.e., the diameter) 
appears to be directly dependent on the richness of the habitat. 
Almost without exception, the lots, with a mean major diam- 
eter of over 10 mm., occur in the most heavily wooded por- 
tions of the limestone or central region; these places are 
usually near the larger hills, which probably increase the rain- 


100 University of Michigan 


fall in their near vicinity. The number of whorls (and the 
altitude) appears to be inversely proportional to the amount 
of exposure to the dry trade-winds; this factor probably acts 
through increase in the rate of evaporation, which would de- 
crease the length of the active periods of the cerions. All of 
the lots, which have a mean number of whorls that approaches | 
12, are from the base of escarpments, usually on the lee side 
of the larger hills. 

Typical C. uva uva probably comes from the hills behind 
Willemstad. This lot (Table XI) happens to give a mean size 
and the mean number of whorls near those of all of the lots 
taken together. Although the change in form, due to the num- 
ber of whorls and the resultant difference in altitude, is very 
conspicuous (Plate XVIII), it is too variable to be of any 
racial importance; in fact, aperture formation appears to be 
hastened by injury. The actual size of the shells seems more 
important, and the extreme lots deserve recognition as ecologi- 
cal forms. The most dwarfed shells may be called form 
diablensis, new (fig. xviili-A2), with the top of Ronde Klip 
(C12b) as the type locality. The largest shells also have the 
heaviest sculpture, and may be included in the form hatoensis, 
new (fig. xviii-F6), with the eastern escarpment of Seroe 
Spelonk, near Landhuis Hato (C11d) as the type locality. 

C. uva uva is mainly restricted to the limestone outcrops, 
but in a few places invades the borders of the central region 
of older rocks. In these localities (Cb6, 10), a rather large 
portion of the shells show a tendency to reduce the sculpture 
(figs. xvili-C6, D5), and even the sculptured shells are usually 
rather slender. Practically smooth shells occur, and have been 
described as C. uva desculptum; the exact type locality of 
this very conspicuous form is unknown, but it is probably 
somewhere around Sint Anna Baai (Cb6b). 


Occasional Papers of the Museum of Zoology 101 


Table XI. Variation in Cerion wva wa from Substations C5c, d44 
A. Variation by Altitude Class 
Whorl Class (by half-whorls) 


Alt. Mean 
Class.) S100) )l0!5) 210) Sse 1220) 12:5) 130) “Whorls 
17.0 al 3 10.4 
17.9 1 5 4 10.6 
18.8 10 19 5 10.9 
19.7 5 49 25 2 iio 
20.6 3 27 48 14 11.4 
21.5 5 49 24 3 11.6 
22.4 2 ah 35 4 11.9 
23.3 +f if 12.3 
24.2 i 2 12.3 
25.1 2 13.0 
Mean 


Altitude 17.4 188 198 20.8 218 22.8 25.1 


B. Variation by Major Diameter Class 
Whorl Class (by half-whorls) 


Maj. Diam. Mean 
Class 10.0 105 11.0 115 12.0 12.5 13.0 Whorls 
8.6 1 4 3 Z 2 11.0 
9.0 3 15 19 12 } 11.4 
9.4 7 37 47 25 6 1 11.5 
9.8 1 8 34 47 26 7 11.4 
10.2 4 14 20 11 2 1 11.5 
10.6 3 4 2 li 11.5 
Mean 


Maj. Diam. 9.2 9.4 9.6 9.6 9.6 Det 9.8 


C. Variation by Major Diameter Index Class 
Whorl Class (by half-whorls) 


Index Mean 
Classe LOO etOloe IO) 9 115 2 TO 2b 13:0) Wihorls 
56 2 10.5 
53 1 5 10 1 10.8 
50 il 11 39 23 S alalaal 
47 8 47 64 14 1 11.3 
44 10 46 42 8 ili ley( 
41 4 18 Uf if iP 
38 1 iL 1 12.5 

Mean 


Index 51 50 48 46 44 43 39 


44 The columns of figures in the central portions of the subtables give 
the number of individuals which fall into each category. 


Cerion uva knipensis, new subspecies 


102 University of Michigan 
Table XII. Mean and Extreme Dimensions of Cerion wa uva 
Comparison of Stations 

Place Nos. Whorls Altitude Index Maj. Diam. 
Clld 57 11.8(10.5-13.0) 22.1(18.8-25.0) 46(42-54) 10.3(9.3-11.3) 
Cl1le 104 11.6(10.5-13.0) 21.0(18.5-24.1) 48(40-60) 10.2(9.1-11.4) 
C6e 77 11.5(10.0-12.5) 21.2(17.6-23.6) 48(41-60) 10.2(8.9-11.2) 
Cl 32 11.1(10.0-12.0) 20.0(16.8-23.4) 51(43-59) 10.2(9.3-11.2) 
Cl2a 53 12.0(11.0-13.0)a 23.1(20.6-25.3) 44(39-51) 10.1(9.4-11.0) 
C4 103 11.6(10.5-13.0) 21.8(18.7-25.2) 46(39-55) 10.1(9.1-11.1) 
C8 60 11.8(10.5-13.0) 22.1(19.1-15.1) 45(40-52) 10.0(9.2-11.2) 
C2 122 11.9(10.0-14.0) 21.8(17.6-26.4) 46(37-60) 9.9(8.2-10.9) 
Cllab 70 11.2(10.0-12.5) 20.0(16.5-23.7)b 48(41-55) 9.9(8.2-10.6) 
C7ab 95 11.3(10.5-12.5) 21.0(18.3-24.0) 47(41-53) 9.8(8.9-10.8) 
C13 51 12.2(10.5-13.5) 22.1(17.0-26.6) 44(37-56) 9.7(8.9-10.7) 
C9 10 11.3(10.5-12.5) 19.6(17.5-21.5) 49(45-59) 9.7(9.2-10.2) 
C6ab 67 11.6(10.0-12.5) 21.3(18.1-24.7) 45(39-52) 9.6(8.6—10.4) 
Cded 370 11.4(10.0-13.0) 20.7(17.0-25.2) 46(38-56) 9.6(8.6—10.8) 
Cdab 110 11.4(10.5-12.5) 20.3(17.9-24.2) 46(40-54) 9.6(8.6-11.2) 
Cb6 52 11.6(10.5-12.5) 21.3(19.38-24.7) 44(37-50) 9.5(8.7-10.1) 
C3 54 11.4(10.5-12.5) 20.0(17.3-23.1) 47(40-55) 9.5(8.4-10.3) 
Cb10 45 11.7(10.5-13.5) 21.3(18.7-25.5) 44(37-52) 9.4(8.4-10.3) 
C10 48 11.3(10.0-12.5) 19.8(16.8-23.6)¢ 47(38-54) 9.2(8.5-10.1) 
C12b 63 11.5(10.5-12.5) 20.1(17.4-22.6) 45(39-52) _9.1(7.8-10.0) 
Total 1643 11.6(10.0-14.0) 21.0(16.5-26.6) 46(37-60) 9.8(7.8—11.4) 

a Bimodal: 11.5 and 12.5; b bimodal: 19.7 and 21.5; ¢ bimodal: 18.8 and 21.5. 


Type locality: (Cb17b) valley between Seroes Palomba and 
Baha Hoendoe, northern Curacao. 
Distribution: Curacao; north of Seroe Grandi; both on 
limestone (C1420), and in the higher hills on outerops of 


the older rocks (Cb16, 17, 20). 
burrowing more deeply in the talus of the older rocks. 


adults collected. 
This subspecies (Plate XIX) has higher whorls than typical 
In the statistical study, the data for each lot were ar- 


Uva. 


Similar in habits to wva, but 


429 


ranged in three tables, with vertical columns for the whorl 
classes and transverse rows to indicate, in the different tables, 
the altitude, major diameter and index classes (cf. Table XI). 
This brought out the fact that, in C. wva uva, the altitude was 
very closely correlated with the number of whorls, but did not 


Occasional Papers of the Museum of Zoology 103 
vary greatly with the diameter. Although considerable indi- 
vidual variation is present, the means show that C. wva knipen- 
sis (Table XIII) quite consistently has a greater altitude for 
the same number of whorls. The only lots of C. wva uva that 
tend to approach them, in this particular, are the very large 
shells from near Landhuis Hato (Clle, d, and Cl12a). 

The typical specimens (fig. xix-F'5) of this subspecies come 
from the wooded valleys of the higher hills, in the region of 
the older rocks. These shells are very large, and have a pecul- 
iar, porcellanous texture, which contrasts rather markedly 
with the chalky surface of most of the shells from the lime- 
stone outcrops. In addition, a few specimens show a slight 
tendency to reduce the sculpture, although most of them are 
quite as heavily sculptured as typical C. uwva wva. 

C. wa knipensis (Table XIII) shows the same type of 
ecological variation discussed in C. uva uva (Table XII). 
Many of the animals mature with a smaller number of whorls 
than do any of the typical subspecies. In addition, there is a 
similar amount of variation in the major diameter. The small- 
est specimens may be called form djerimensis, new (fig. 
xix—Al), with the top of the shore cliffs near Plaja Djerimi 
(C17c) as the type locality. j 


Table XIII. Mean and Extreme Dimensions of Cerion wa knipensis 
Comparison of Stations 


Place Nos. Whorls Altitude Index Maj. Diam. 
C18 24 11.2(10.5-12.0) 21.3(18.3+23.6) 49(44-55) 10.5(9.5-11.2) 
Cb17, 20 72 11.9(11.0-13.0) 24.0(20.5-27.8)a 43(34-50) 10.3(9.3-11.2) 
Cb16 16 11.4(11.0-12.5) 22.8(20.9-26.1) 45(37-49) 10.1(9.6—10.6) 
C20 31 11.5(10.5-12.5) 22.5(19.8-25.6) 45(37-51) 10.0(9.1-10.7) 
C16 34 11.2(10.5-12.5) 21.7(19.4-25.6) 46(39-51) 10.0(9.0—-10.5) 
Cl7a 55 11.1(10.5-12.5) 21.2(18.8-24.8) 47(42-53) 9.9(9.1-11.0) 
Olds 17a 81 11.1(10.0-12.5) 21.2(18.3-24.2) 46(39-53) 9.7(8.7—-10.6) 
C19 41 10.5( 9.5-11.5) 20.2(17.6-22.7) 48(44-55) 9.6(8.8—10.5) 
C14 54 10.8( 9.5-12.0) 20.0(17.2-24.3) 47(37-53) 9.4(8.6—10.6) 
C17be 21 10.5( 9.5-11.5) 19.7(17.2-24.3) 46(39-53) 9.1(8.1-10.1) 
Total 429 11.1( 9.5-13.0) 21.5(17.2-27.8) 46(34-55) 9.9(8.1-11.2) 


aBimodal: 22.4 and 25.1. 


104 University of Michigan 


Cerion uva arubanum, new subspecies 


Cerion uva Smith (1898; Proc. Mal. Soe., III, 114); the first Aruba 
record; collected by Hartert. 

Type locality: (A2c) Baranca Alto, Aruba. 

Distribution: Aruba; living shells only found in a colony at 
the type locality and in another just north of Perkietenboseh 
(A2c,A5a); subfossil throughout the limestone portions of 
the island. Similar in habits to wva. 322 adults collected. 

Although considerable individual variation (Plate XX) 
occurs, the last whorl near the aperture tends to jut out tan- 
gentially in this subspecies, so that the palatal wall of the 
peristome usually projects out from the preceding whorls 
to a greater extent than in C. uva uva and knipensis. As a 
result of this, the umbilicus is usually larger and more open 
in the specimens from Aruba. In making the measurements 
of the major diameter in the two subspecies from Curacao, 
the calipers seldom touched the aperture, while in C. uva aru- 
banum, the palatal wall almost always interfered. In all eases, 
the major diameter was taken exclusive of the aperture, so 
the true greatest width of C. wa arubanum is slightly larger 
than the data in Table XIV would indicate. In other par- 
ticulars, this form is similar to the smaller and more slender 
lots of C. wa uva, although the whorls tend to be slightly 
lower than in any of the Curacao lots except those from 
station C13. 

The peculiar restriction of the living cerions on Aruba to 
two isolated colonies is very puzzling, especially since this 
species very evidently was almost universally distributed on 
the island in former times. Subfossil shells occur almost 
everywhere on the limestone, and also are cemented into blocks 
of phosphate near Culebra (A1). It does not seem possible 
that this former distribution has been reduced by the extensive 
cultivation of aloes, as living shells actually occur on this plant 
in station A4e, while they are absent from the large, limestone 
plateaus east of Savaneta and Sint Nicolaas, and these hills 
appear to be mainly undisturbed by man. Although Aruba 
has a more arid climate than the other islands and appears 


Occasional Papers of the Museum of Zoology 105 
to be more subject to pronounced dry periods, it is rather diffi- 
cult to understand how such a resistant species as Cerion uva 
could have been partially exterminated by these factors. In 
addition, the living colonies are not in the most heavily wooded 
places. _ 

The specimens from the two colonies differ slightly from 
each other in dimensions, and one lot is lighter in color than 
the other. However, these divergences hardly seem worthy 
of rank as separate forms. Some specimens show a slight 


approach towards C. wa desculptum. 


Cerion uva bonairensis, new subspecies 


Type locality: (B5) Porta Spano, Bonaire. 

Distribution : Bonaire ; everywhere on limestone (B1 to B9), 
and in the richer habitats on the older rocks (Bb3, 7, 9). 
Klein-Bonaire: everywhere (K1). 348 adults collected. 


Table XIV. Mean and Extreme Dimensions of Cerion wa arubanum 
and bonairensis 


Comparison of Stations 


Place Nos Whorls Altitude Index Maj. Diam. 
A2e 200 11.9(11.0-13.5) 21.6(18.7-25.9) 44(37-51) 9.4(8.3-10.3) 
Ada 122 12.2(11.0-14.0) 22.5(20.0-27.4) 43(36-50) 9.6(8.7—10.8) 
Totals, 

Aruba 322 12.0(11.0-14.0) 22.0(18.7-27.4) 43(36-51) 9.5(8.3-10.8) 
Bb3 12 11.7(11.0-12.5) 24.1(22.2-25.8) 43(40-48) 10.3(9.9-10.7) 
B5 39 11.5(10.5-12.5) 22.2(19.6-26.8) 44(39-53) 10.3(8.9-11.0) 
B8 58 11.3(10.5-12.5) 21.8(19.7-24.4) 45(38-49) 9.9(9.0—-10.8) 
B7 51 11.5(10.5-12.0) 22.4(19.6-25.4) 44(39-50)- 9.8(9.0-10.9) 
Bb7 20 11.5(10.5-13.0) 22.3(19.7-25.4) 44(39-50) 9.7(9.0-10.2) 
B4 17) =11.4(10.5-12.5) 21.9(19.2-24.1) 44(37-51) 9.7(8.4-10.3) 
K1 28 11.2( 9.5-12.0) 21.2(16.0-24.2) 46(41-61) 9.7(8.9-10.7) 
B3 44 10.9( 9.5-12.0) 21.0(17.1-23.1) 47(41-55) 9.7(8.9-10.5) 
B6 6 10.8(10.5-11.5) 19.7(19.2-20.2) 49(47-51) 9.6(9.1—-10.1) 
B9, Bb9 28 11.1(10.0-12.0) 20.4(18.2-23.9) 45(38-53) 9.1(8.2—10.2) 
Bl 31 10.9(10.0-11.5) 20.0(17.8-21.2) 46(41-52) 9.1(8.4— 9.6) 
B2 9 10.7(10.0-12.0) 19.7(17.2-23.1) 47(42-51) 9.1(8.4—10.1) 
Totals, 

Bonaire 343 11.2( 9.5-13.0) 21.6(16.0-26.8) 45(37-61) 9.7(8.2-11.0) 


106 University of Michigan 


In this subspecies (Plate X XI), as in arubanum, the palatal 
wall of the peristome projects out markedly from the preced- 
ing whorls. However, the aperture is usually broader, instead 
of twisted tangentially, so that the umbilicus is smaller, and 
more like that of the typical subspecies. Also, C. uva bonmr- 
ensis (Table XIV) has high whorls similar to knipensis, and 
the cleaned surface of the shell resembles dead-white enamel 
while that of the other species is chalky-white or porcellanous. 
As in the subspecies from Curacao, the Bonaire lots vary in 
size with the richness of the locality. Typical bonairensis 
(fig. xxi-F'6) includes the larger shells of the series, while the 
smaller lots may be called form kralendijki, new (fig. xxi-A2), 
with the recent limestone just south of Kralendijk (B1) as 
the type locality. Although the cerions of Bonaire invade the 
central regions of older rocks to a considerable extent, the form 
desculptum appears to be quite absent. 


INTRODUCED AND DOUBTFUL SPECIES 


Outside of the hofje of Campo Knip, and a few borders near 
the highways, no detailed examination of the cultivated ground 
was made. A more thorough study of the irrigated fields and 
gardens would probably add considerably to the following list. 
Some of the coco plantations and orchards are several acres in 
extent, and the borders of the better watered ones have de- 
veloped considerable leaf mould. However, such introduced 
" species have no bearing on the zodgeographical affinities of the 
islands and, while interesting, would certainly add very little 
to our knowledge of the natural fauna. 


TRUNCATELLIDAE 
Truncatella bilabiata (Pfeiffer) 

The last 1144 whorls of a bleached shell, from the hofje of 
Campo Klein Piscadera (Cb7). This belongs to the almost 
smooth form of the species, and the growth-ribs are only rep- 
resented by buttresses near the suture. 


. HELICIDAE 


Thysanophora crinita arubana H. Burrington Baker 
One dead, bleached specimen, from the hofje of Klein Pisea- 


= 


Occasional Papers of the Museum of Zoology 107 


dera (Cb7), is probably this form, although its condition does 
not permit of its certain identification. In this connection, it 
is at least interesting that the only amphibian collected in 
Curacao, Pleurodema brachyops (Cope), is said by the in- 
habitants to have been introduced from Aruba in sand for 
construction. 


Pleurodonte lima (Férussac) 


Heliz lima Férussae (1821; Tabl. Syst., 32; Hist., fig. xlvi-l, 2). HZ. 
lima Pfr. (1848; Mon. Helic. Viv., I, 266); exclusive of var. B. Pleuro- 
donte lima Pilsbry (1889; Man. Conch., V, 58); exclusive of Otala 
asperula, in synonymy, and the reference to Curagao. 


As discussed under the next species, there is no reason for 
the citation of P. lima from Curaeao. 


Pleurodonte incerta (Férussac) 

Helix lima ‘‘variété et jeune age’’ Férussac (Hist., fig. xlvi, A-44, 5). 
Helix incerta Férussac (Hist., fig. ev-2). Otala asperula Beck (1837; 
Index, 36) ; founded on ‘‘H. Hgen. lima, var. notabilior Fér.; F. H., 
xlv, A, 4-5.’’ Helix lima, var. B Pfr. (1848; Mon. Helic. Viv., I, 266). 
H. lima var. notabilis ‘‘ Fér.’’ Pfr. (l. c.); in synonymy. H. incerta var. 
notabilis Albers-Martens (1860; Die Heliceen, 144). Plewrodonte incerta 
Pilsbry (1889; Man. Conch., V, 57-58). Plewrodonte incerta Vernhout 
(1914; Notes Leyden Mus., 179). P. lima Vernhout (l.c.). 


In the Histoire, Férussae illustrated his typical form of 
H. lima on Plate 46, but also figured an unnamed ‘‘variety’’ 
on Plate 46A. Beck founded his Otala asperula on this ‘‘nota- 
bilior’’ variety,*® and added ‘‘I. Curacao’’ as the type locality 
of his new species. Pfeiffer placed Beck’s species, along with 
““ HH. lima var. notabilis Fér.’’ (the first use of the latter name), 
in the synonymy of his H. lima, var. 8, and quoted Curacao 
as the locality. On the other hand, Albers-Martens regarded 
notabilis as a variety of H. incerta, and so, very naturally, 
included Curacao as the habitat. Pilsbry followed Albers- 
Martens in the disposition of notabilis, but copied Pfeiffer in 
the inclusion of Otala asperula, and its locality, under H. lima. 


45 xlvy, A is obviously a misprint for Plate 46A. 


108 University of Michigan 


Thus all of the Curacao citations of either of these two species, 
and Vernhout’s quotation of both of them, may be traced back 
to Beck’s exceedingly dubious record. Finally, if Férussae’s 
variety requires a name, it must be called Plewrodonte incerta 
asperula (Beck) ; this form is now known to occur in Porto 
Rico. 

BULIMULIDAE 


Drymaeus multilineatus (Say) 


Bulimus multilineatus Say (1825; Jour. A. N. 8. P., V, 119). Bulimus 
torallyi, var. B. sisalensis Bland (1868; Amer. J. Conch., 192); first 
Curacao record. 


Type locality: southern part of east Florida. 

Distribution: southern Florida and Yucatan to northern 
South America. 

This species has been quoted from Curacao by several 
authors, and two lots in the A. N. S. P. (no. 25896 and 25900; 
Swift Collection, collected by Raven) are certainly nothing 
else (cf. Pilsbry; 1899; Man. Conch., XII; p. 29 and fig. 
xi-32). In 1920, I collected two dead bleached specimens 
on the Schaarlo (C5c), at the edge of Willemstad, but no addi- 
tional specimens were obtained in 1922. It may still occur 
in the gardens in Willemstad, or on some of the irrigated 
plantations. This species thrives in cultivated places, and I 
have no doubt that it has been introduced into many parts of 
its present range. It does not appear to be established, under 
natural conditions, on Curae¢ao. 


LTigwus virgineus (Lin.) 

This Hispanolan species has been listed from Curacao by 
Vernhout (1914; Notes Leyden Mus., 179, 180). As I did 
not obtain it, I can only add that, as yet, it does not appear 
to have escaped from cultivation. 


ACHATINIDAE 
Opeas micra (D’Orbigny ) 


Stenogyra octonoides Gibbons (1879; J. of Conch., II, 136); first ree- 
ord from Curacao. 


Occasional Papers of the Museum of Zoology 109 


Two dead specimens were obtained from leaf mould along 
the road at Campo Klein Piscadera (Cb7). This species has 
been widely distributed by commerce. 


ZOOGEOGRAPHICAL AFFINITIES OF THE MoLLUSCAN FAUNA 


In the study of the terrestrial, molluscan fauna of these 
islands, five, salient characteristics appear. (1) There is a very 
high percentage of endemism. (2) Sublittoral, limestone-loving 
species preponderate. (3) Typically South American groups 
are completely absent. (4) Several Antillean groups form 
conspicuous components of the total population. (5) Neverthe- 
less, the closest affinities shown by any of the individual species 
are with forms from northern South America. 

Two genera (Stoastomops and Cistulops), three subgenera 
(Bonairea, Neosubulina s. s., and Cerion s. s.), and a section 
(Tudora s.s.) are, as far as known, endemic to the Dutch 
Leeward Islands. Only three of the well-established species, 
Drymaeus virgulatus, Thysanophora crinita and Oxystyla 
maracaibensis, are known to extend their range beyond the 
islands. The first of these is a species of Porto Rico and the 
northern Lesser Antilles. Although it is certainly widespread 
in the undisturbed portion of the Dutch Leeward Islands, its 
remarkably discontinuous distribution, as already indicated, 
arouses the suspicion that its dissemination may be due to the 
agency of man. The last two are species of northern South 
America, and only reach Aruba (although 7. crinita was 
found in ruderal conditions on Curacao); the Oxystyla ap- 
pears to be entirely subfossil. 

The endemism of the fauna of Bonaire, Klein-Bonaire and 
Curacao is readily accounted for on the basis of their geo- 
graphic isolation. As already described, the depth of ocean 
between these islands and the mainland indicates a separation 
at least as remote as that between South America and the 
northern Lesser Antilles. Aruba, on the other hand, lies in 
quite shallow water, and it is rather remarkable that its fauna 
does not contain an even larger proportion of South American 
invaders. 


110 University of Michigan 


Practically all of the molluses of the islands show a marked 
preference for limestone rock, and most of them do not occur 
in localities where the soil is non-caleareous. Drymaeus virgu- 
latus, Succinea gyrata and Guppya molengraaffi are the con- 
spicuous exceptions to this general rule. The first two appear 
to be mainly dependent on the development of trees and brush, 
while the last was only found near the summit of Sint Chris- 
toffelberg (Cb20). Although several species invade the more 
heavily wooded portions in the higher hills of the older rocks, 
they are much rarer in these places than on the limestone. 
The deep aestivation of Zudora fossor and the loss of sculp- 
ture in Cerion uva have already been correlated with this 
species. 

In addition, the fauna is characteristic of coastal conditions. 
Cerion, especially, is usually limited, throughout its distribu- 
tion, to the near vicinity of the seashore. The highland rain- 
forest elements of both the Antilles and South America have 
not obtained a footing on the present islands, although Guppya 
molengraaffi appears to be limited to the region where such 
conditions are most nearly approached. In places where such 
a mesophytie fauna is present, its more hardy members may 
invade the coastal region, but in the Dutch Leeward Islands 
the opposite is true, and even the cerions reach to the highest 
altitudes (1,200 feet). 

As a result of these ecological limitations, the fauna is only 
comparable to that of similar regions in the Antilles and on 
the mainland. This type of habitat has been quite thoroughly 
studied in the former, but, in South America, collectors have 
directed most of their attention to the richer inland forests. 
A detailed study of the caleareous portion of the shore zone of 
northern South America (cf. W. Sievers; 1896; 1. c., pl. X) 
might add considerably to our knowledge of the Antillean 
elements in this region. On the basis of the present data, it 
can only be stated that Aruba, as well as the other Dutch Lee- 
ward Islands, is markedly isolated ecologically from the known 
portions of the mainland. The similarity of the fauna of 
Aruba and that of the other islands and its lack of resemblance 


Occasional Papers of the Museum of Zoology 111 


to the known South American fauna, can only be explained 
on this basis. 

As already indicated, one of the most notable features of 
this island fauna is the total absence of typically South Amer- 
ican groups, such as the Streptaxidae and the Strophocheili- 
nae, and the paucity of others, such as the Achatinidae and 
Bulimulidae. In fact, not a single, characteristically South 
American genus reaches these islands, although four (Dry- 
maeus, Oxystyla, Thysanophora and Guppya) might be con- 
sidered as widely distributed, tropical or subtropical groups 
which penetrate South America to a considerable extent. 

On the other hand, Cerion is practically limited to the 
northern Antilles and Florida, although a single species, Cerion 
antonu (Kuester), has been deseribed from Guiana. Micro- 
ceramus is another Antillean genus, which reaches the main- 
land of North America but is apparently lacking from the 
southern Antilles and South America. Stoastomops is, I be- 
heve, most closely related to Stoastoma from Jamaica, while 
Cistulops appears to have distant affinities with Troschelvin- 
dex from central Cuba. The Chondropominae are also a char- 
acteristically Antillean group, although they reach the main- 
land in many places around the Caribbean and the Gulf of 
Mexico. In addition, the two freshwater species, Potamopyr- 
gus parvulus and Planorbis pallidus, are widely distributed 
in the Antilles, although they are represented by closely re- 
lated species in northern South America. 

Although the molluscan fauna of these islands is thus 
mainly Antillean in its general affinities, some of the individual 
groups are most closely related to those of northern South 
America. For example, Tudora rupis, T. muskusi and T. 
aurantia are placed in the section Tudorata, along with Tudora 
plicatula from Venezuela. Also, the genus Neosubulina is 
known only from the Dutch Leeward Islands and northern 
South America, while Brachypodella raveni and gibbonsi be- 
long to the mainland group of their genus. Finally, Oxystyla 
maracaibensis imitator and Thysanophora crinita have already 
been listed as South American species that also occur in Aruba. 


112 University of Michigan 


These peculiarities of distribution may be explained by the 
hypothesis that the molluscan species of the Dutch Leeward 
Islands are considerably changed remnants of the ancient 
Antillean fauna of Archiguiana (cf. Von Ihering, 1907; 
Archhelenis and Archinotis, p. 111). On the mainland, this 
fauna is largely replaced by Brazilian and northern, conti- 
nental elements, although it may still be represented by 
Brachypodella, various Pomatiasidae, probably such groups 
as Sericea, Analcadia and Tamsiana among the Helicinidae, 
and the Lesser Antillean group of Amphicyclotus among the 
Cyclophoridae. The Oleacinid group Ravenia from Los 
Roques is another Antillean group of this same general region. 
In addition, there are a number of doubtful species, such as 
Eutrochatella semilirata, Helicina kienert and Chondropoma 
subauriculatum, described as from Venezuela (cf. no. 137 of 
this series), and Cerion antonu, cited above, which arouse the 
suspicion that a more thorough study of the northern coast 
of South America may bring a greater number of these 
‘*relies’’ to light. 

However, it is somewhat doubtful if the Dutch Leeward 
Islands could have retained their species as the direct and con- 
tinuous descendants of the faunas of such a land mass. The 
present altitude of such late Tertiary coral formations as 
the cap of the Tafelberg of Sint Hyronimus (C20) seem to 
indicate a possibility that the islands were completely sub- 
merged in comparatively recent times. Much of the present 
fauna of the Dutch Leeward Islands and the presence of An- 
tillean elements in northern South America is also explicable 
on the basis that these are all comparatively recent arrivals 
by drift or other occasional means of dispersal, and that some 
of them are only able to gain a foothold in places where they 
do not come into competition with the continental species. 
This is especially true of the fresh water species; their pres- 
ence in artificial ponds and reservoirs certainly indicate ex- 
traordinary means of dispersal, such as carriage by birds. 
But, such endemic groups as Cistulops, Stoastomops, Bonairea 
and Cerion s.s. can scarcely be explained by this alternative 


Occasional Papers of the Museum of Zoology 113 


hypothesis. The question must be left in abeyance, at least 
until a thorough examination of the remainder of the Leeward 
Islands and the northern coast of South America has been 
made. 


DISTRIBUTION WITHIN THE ISLANDS 


As most of the species of the Dutch Leeward Islands are 
almost entirely limited to the limestone rim, breaks in this 
establish quite definite barriers to intermigration. Especially 
on the island of Curacao, the sunken valleys permit penetra- 
tion by arms of the sea, so that the rim is cut up into a series 
of quite isolated ridges. As a result, the facies of certain 
species of the land shells changes slightly at each ‘‘baai’’ or 
‘‘lagoen.’’? After several successive breaks, the species of 
Tudora, especially, often change so much that there is no inter- 
gradation between lots from the colonies so separated. It is 
on such changes that the various subspecies are based. 

The largest island, Curacao, can be divided into three, quite 
distinct, faunal areas, which appear to have been populated 
from central hills, which must have been separate islands 
during periods of higher strand-line. Aruba, on the other 
hand, is almost a unit, although Spaansch Lagoen roughly 
coincides with a slight break in the mollusean fauna (see Plate 
I, map of Aruba). Bonaire and Klein-Bonaire together form 
a similar unit. 

The most southern area of Curacao centers around the 
Tafelberg of Santa Barbara (C2). Tudora rupis rupis, T. 
pilsbryi and Brachypodella raven sanctaebarbarae were only 
found at the base of the northern and western escarpments 
of this mesa. The first species is also represented by the sub- 
species newportensis near New Port (C1), although the re- 
mainder of the fauna of the shore zone is more like that of 
central Curacao. In addition, Cistulops raveni appears to be 
absent from southern Curagao. 

The central area (C1, 3-12) may be considered to center 
around Ronde Klip (C12) and the Hato ridge (C11). It is 
characterized by the absence of the section Tudorata, and by 


114 University of Michigan 


the presence of Tudora megacheilos (4 subspecies) and 
Brachypodella raveni raven. 

The northern area (C13-20) was probably populated from 
the Sint Christoffel complex, which undoubtedly was formerly 
fringed or covered by a limestone reef, such as is still present 
on the Tafelberg of Sint Hyronimus (C20). This region is 
differentiated by Tudora muskusi (3 subspecies), Guppya 
molengraaffi, 3 subspecies of Tudora fossor, Brachypodella 
ravemi knipensis and Cerion uva knipensis. 

Neosubulina gloynti and Microceramus bonairensis cwracoa- 
nus occur throughout Curacao. The northern and central 
areas agree in the occurrence of Cistulops ravem ravent, while 
the central and southern possess Cerion uva uva in common. 
The presence of the section Tudorata, and the resemblance of 
the subspecies of Brachypodella and the species T'udora fossor 
and pilsbryi, appear to relate the northern and southern areas 
more closely to each other than to the central portion. The 
first two peculiarities of these terminal regions also seem to 
indicate affinities with the island of Bonaire. 

The island of Aruba has, for peculiar forms, Neosubulina 
scopulorum, Thysanophora vanatta, Cistulops ravent arubana, 
Brachypodella raveni arubana and the form sinistrorsa, Micro- 
ceramus bonairensis arubanus, Cerion uva arubanum, two sub- 
species of T'udora fossor, and the two South American forms, 
Thysanophora crinita arubana and Oxystyla maracaibensis 
imitator. In general, the mollusecan fauna of this island is 
most closely related to that of northern Curacao, with which 
it possesses Tudora fossor in common; the subspecies of 
Brachypodella raveni are also similar. However, Cerion uva 
arubanum appears most closely related with C. wea wva from 
central Curacao and Tudorata is similarly absent in both 
regions. 

Bonaire and Klein-Bonaire together form the most distinet 
of all of the areas, with the genus Stoastomops, the subgenus 
Bonairea, Tudora aurantia, Neosubulina harterti (the most 
remote of the three species), Brachypodella gibbonsi, Micro- 
ceramus bonairensis bonairensis, Cerion uva bonairensis, and 


Occasional Papers of the Museum of Zoology 115 


the absence of Cistulops and Tudora s.s. as points of differ- 
ence. In this connection, it is at least interesting that the 
direction of the ocean current is from Bonaire towards 
Curacao, and from the latter to Aruba; it is considerably 
stronger between the two more western islands. 

Bonaire and Curacao agree in the absence of Thysanophora 
and Oxystyla, and in the presence of the section Tudorata and 
the species Succinea gyrata. 

These differences in molluscan fauna appear to be almost 
purely geographical ; that is, they are coincident with degrees 
of present or former isolation. The greatest ecological dif- 
ferentiation is between the abundance of individuals and spe- 
cies on the caleareous substrata and their paucity or absence 
on the non-caleareous soil. Except for certain minor edaphic 
phases, represented by the forms of Cerion wva, the formation 
on all of the islands is practically identical, as are also most of 
the factors of the environment. In other words, the formation 
content (the mollusean association or society) changes with 
the area, although its general aspect remains the same. 

As an illustration of this, the habits of Tudora on Curacao 
may be cited. In northern and southern Curacao, the species 
of Tudora s.s. (7. fossor and T. pilsbryt) are almost purely 
terrestrial, while the species of Tudorata (7. rwpis and T. 
muskust) are markedly subarboreal. In central Curacao, 
Tudorata is absent, but 7. (s.s.) megacheilos differs from its 
closest relatives in a well-developed tendency to climb the 
trees and brush, so that it practically occupies both the ground 
and lower arboreal strata. Judora fossor arubana also shows 
this tendency, although it is much less prominent. 

In this connection, it may be remarked that Cerion wva is 
also an adept climber during the rains, but appears to migrate 
down to fasten itself near the base of the trees and brush dur- 
ing the dry periods. Drymaeus virgulatus, on the other hand, 
commonly glues itself to the highest branches; this is perhaps 
related with its restriction to the more protected stations. 
Finally, the subarboreal species of Tudora wander up the trees 
in the rainy weather, but have not developed the opposite 


116 University of Michigan 


reaction. However, with the operculum closed, they are unable 
to attach themselves firmly to the branches and so are usually 
shaken out in a few days by the wind. Nevertheless, specimens 
often remain in hollows and erevices of the bark, although 
their survival in such places appears to be comparatively rare. 
Cerion uva appears more resistant to dessication, but the secre- 
tive habits of Tudora enable it to withstand an equal amount 
of aridity; the exposed cerions are dwarfed in the most arid 
localities, but size in Tudora appears to have little or no 
edaphic significance. 

In conclusion, the most remarkable case of parallel fune- 
tions, in different ‘‘cenoses’’ but in the same ‘‘mores’’ is 
furnished by Cistulops raveni (Curacao and Aruba) and 
Tudora maculata (Bonaire and Klein-Bonaire). These two 
distantly related species are almost identical in habits and 
must occupy practically the same position in their respective 
societies. Although they differ in radula, operculum and 
peristome, they resemble each other so closely in form, size, 
color and sculpture, that, when I first found 7. maculata, I 
spent some time in a search for specimens with well-developed 
apertures, under the impression that I was collecting im- 
mature specimens of C. raveni or some closely related species! 
As already indicated, Thomas Bland probably neglected to 
publish his ‘‘ Cistula maculata’’ for the very same reason. 


ADDENDA 
Since the preceding paper was finished, Dr. H. A. Pilsbry 
has worked up the Pupillidae collected, and is publishing their 
descriptions in the Proc. Acad. Nat. Sci. Philadelphia (1924). 
As indicated below, he recognizes three species of Gastrocepta, 
instead of G. longurio, and one species of Pupoides. 


Gastrocopta curacoana Pilsbry 
Widely distributed on Curacao (C1-6, 11-15, 17), Aruba 
(A2, 4, 5, 8), Bonaire (B1-5, 8) and Klein-Bonaire (K1). 
Type locality: Fort Nassau, Curacao (C5d); new name for 
Pupa longurio Crosse (1872), not Moquin-Tandon (1855). 


Occasional Papers of the Museum of Zoology 117 


Gastrocopta octonaria Pilsbry 
Widely distributed on Curacao (C1, 2, 5, 6, 10-15, 17, 18), 
Aruba (A2-4, 8) and Bonaire (B3, 4, 8). Type locality: Fort 
Nassau, Curacao (C5dd). 


Gastrocopta barbadensis hojeda Pilsbry 
Curacao: Tafelberg of Santa Barbara and north of Playa 
Abau (C2, 18). Aruba: Rooi Frances and Seroe Canashito 
(A3, 4). Type locality: base of western escarpment of the 
Tafelberg of Santa Barbara, Curacao (C2b). 


Pupoides marginatus nitidulus (Pfr.) 

Distribution in the islands as indicated under Pupoides sp.? 
in the main body of this paper (C3-6, 12, 17; A3—5, 8; B2, 4, 
8; K1). This is a widely distributed, mainly West Indian 
species. According to Dr. Pilsbry, Pupoides simoni (Jous- 
seaume) is probably a synonym. G. barbadensis and P. 
marginatus add two more to the list of non-endemic species 
of the islands. 


118 University of Michigan 


PLATE I 
FIGURE 1 
1—A. New Port (C1). G. Salinja Sint Marie. 
2-B. Tafelberg, Santa Barbara (C2). 10-H. Seroe Boca (C10). 
C. Manzalienja arroyo (Cb2). I. Sint Joris Baai. 


D. Tank, Campo Wilhelmina (Ce2). K. Hill (Cb10). 
11-L. Seroe Markita (Clla). 


M. Ridge north of Ronde Klip (C11b). 

N. Seroe Papaja (Clic). 

O. Near Landhuis Hato (Cll1d). 
12—-P. Ronde Klip (C12). 


E. Spaansche Baai; Spaansch Water. 
3—-F. Hill near Fort Beekenburg (C3). 
G. Caracas Baai. 
4-H. Seroe Mansigna (C4). 


I. Lagoen Jan Tiel. 
5-K. Seroe Spanjo (C5a). 13-R. Seroe Largoe (C13). 
se S. Sint Marie Spring (Ccl13). 
ee ee T. Kerk Sint Willebrordus 
M. Schaarlo (C5c). A : 
N ce area U. Kaap Sint Marie. 
. Fort Nassau (C5d). : 
O. Wi 14-V. Seroe Grandi (C14). 
. Willemstad. : : 
P. Sint Anna Baai. hee 2 
RB. ‘chattecat 15—X. Seroe di Boca (C15). 
Aa pe " Y. Sint Martha Baai. 
—S. Seroe Quinta, Overzijde (C6a). 
: 16-Z. Seroe Baha So (C16). 
bata gi ane a re A. Sint Kruis Baai. 
= st A pons Gee): B. Hill Campo Sint Kruis (Cb16). 
Sera eee 17-C. Seroe Djerimi (C17a). 
W. Piscadera Baai. Knip Baai (C17b) 
X. Campo Marchena (Cb6a). Hae ‘Mee Cle). 
Y. Hill, Campo Blenheim (Cb6b). 


7-Z. Jack Evertzsberg (C7a). . Landhuis Knip (Cb17e). 


A. Sint Michielsberg (C7b). . Hofje, Campo Knip(Cb174d). 

B. Sint Michiel Baai; Salinja. , Pond,Campo Lapean(uaia 

C. Campo Klein-Piscadera (Cb7). 18-K. Hill (C18). 

—D. Seroe Spreit (C8). 19-M. Westpunt (C19). 

E. Seroe Popchie (C9). 20-N. Tafelberg, Sint Hyronimus (C20). 
F. Bullen Baai. P. Sint Christoffelberg (Cb20). 


D 
E 
F. Plaja Djerimi (C17b). 
af 
iE 
L 


FIGURE 3 
A. Culebra (A1). N. Seroe Canashito (A4). 
B. Sint Nicolaas. O. Near Perkietenboseh (A5a). 
C. Boca Grande, and sand dunes. P. Southeast of Oranjestad (A5b). 
D. Near Butucoe (A2a). R. Oranjestad. 
E. Spur of Seroe Pretoe (A2b). S. Limestone near Tanki Schipau (A6). 
F. Baranca Alto (A2c). T. Bubali. 
G. Isla (A2ce). U. Kerk Sint Anna. 
H. Fontein (Ac2). V. Shore near Malmok (A7a). 
I. Savaneta. W. Seroe Annaboei (A7b). 
J. Seroe Jamanota. X. Seroe Hudishibana (A7c). 
K. Rooi Taki (A2c). Y. Campo Westpunt. 
L. Rooi Frances (A3). Z. Limestone near Boedoei (A8). 


M. Spaansch Lagoen. 


OLLUSCS PLATE I 


ere penpmemsmene SF 


CURACAO 


11) omen 
i 
\ 


SKM 


SMILES 


Fic. 1. Map of Curacao, adapted from Netherlands Government 1 /20000 topographic maps. 


‘ny; 


(oe nn ne 


ntour intervals 50 meters. The stippling indicates the approximate extent of the limestone. 


Fic. 2. Map of Klein-Curagao, reduced from U. S. Hydrographic Office chart, number 2154. 
Seale as in fig. 1. 


Seneca 


_ Fic. 3. Map of Aruba. Source, contour interval, scale, and stippling as in fig. 1. 


—— = 
See 


120 University of Michigan 
PLATE II 
FIGURE 4 
A. Pekelmeer, Lacre Punt. M. Porta Spafio (Bd). 
B. South of Kralendijk (B1). N. Traai Montagne (Bb5). 
C. Pos Baca (Bel). O. Campo Bolivia. 
D. Pos Frances. P. Fontein (north of B6). 
E. Kralendijk. R. Punta Blanco (B7). 
F. Along shore near Campo Hato 8S. West of Punta Blanco (Bb7). 
(B2). T. Seroe Wassau (B8). 
G. Southeastern escarpment, Mon-_ U. Salinja Goto. 
tagne (B3a). V. Seroe Grandi (B9). 


HAH 


. Kibra di Montagne (B2). 
. Seroe Grandi (Bb3). 


Rincon. 


- Wooded valley (B4). 
. Seroe Largoe or Montagne. 


. Seroe Brandaris (Bb9). 
. Playa Foenshi. 

. Klein-Bonaire (K1). 
708 


de 
(Kel). 


Cas, Klein-Bonaire 


Mo.uuscs PLATE II 


5 KM 


__ SMILES 


Fig. 4. Map of Bonaire and Klein-Bonaire. Source, contour interval, and stippling 
as in fig. 1. 


122 University of Michigan 


PLATE III 


Fic. 5 (top). Southeast from Jack Evertzsberg, southwestern Curacao. 
In the foreground is the seaward slope of the older limestone (C7a); in 
the background are the similar monadnocks of Veerisberg, Seroe Pretoe 
(C6c; note the fossil beaches), Seroe Domi (C6a), and Fort Nassau 
(C5d) ; while in the far distance, to the right of the center, is the Tafel- 
berg of Santa Barbara (C2). A bit of the Schottegat, and the lower 
inland country, are visible between Jack Evertzsberg and Veerisberg. 

Fic. 6 (middle). South from Landhuis Knip, northwestern Curacao. 
In the foreground is one of the richer, open valleys of northern Curacao; 
the dark masses of vegetation are mainly divi-divi trees, while the inter- 
jacent portions are choked with acacias and Opuntia. To the left of the 
background are the seaward slopes of Seroes Palomba and Bientoe (Cre- 
taceous outcrops), separated by a hidden valley (Cb17a). Farther back 
to the right is the limestone mesa of Seroe Djerimi (Cl17a). 

Fic. 7 (lowest). The northern escarpment of the Schaarlo (C5c), south- 
western Curacao. The foreground shows a protected portion of the recent 
limestone, with a rather dense growth of crotons and acacias. The more 
barren crest of the Schaarlo is at the extreme right. In the left back- 
ground is a bit of the Schottegat with the rounded hills of the inland 
country and the hofjes of the richer valleys; the Tafelberg of Santa 
Barbara (C2) is also visible. 


PLATE IIT 


Mo.Luuscs 


124 University of Michigan 


PLATE IV 


Fic. 8 (top). Eastern escarpment of Seroe Canashito (A4), Aruba. 
In the foreground are the slopes of the older rocks, planted with aloes, 
and with a Cereus fence at the base. In the center are the high, exca- 
vated, ancient limestone cliffs of Seroe Canashito, with the separated 
blocks that are characteristic of these escarpments. The cacti on the 
summit are 15 to 20 feet in height. 

Fie. 9 (middle). North towards Kibra di Montagne (B3b), Bonaire. 
Another view, taken from the same locality as the next. One of the 
better wooded valleys in the limestone. 

Fic. 10 (lowest). Southeastern escarpment of Montagne (B3a), 
Bonaire. These low, broken cliffs of the older limestone are rather well 
wooded. This and figure 8 are quite representative of the main escarp- 
ments of the western limestone rims on all of the islands. 


Monuuscs PLATE IV 


126 University of Michigan 


PLATE V 


Fic. 11. Timberline at Fontein (Ac2), northeastern side of Aruba. 
In the foreground and at the right are the very barren regions of the 
exposed, northeastern shores (compare C10, C20, and A7). The wind- 
distorted thornbushes depend on the small streams; a few tops of coco- 
palms locate the hofje of Fontein. In the left background is a small 
sand dune. 

Fic. 12. Near Perkietenboseh (A5a), southwestern side of Aruba. 
This shows the zone of natural vegetation near the summit of the low 
shore cliffs, on the recent limestone. Much of the ground vegetation con- 
sists of Jatropha urens and Opuntia. The divi-divi trees are character- 
istically distorted by the wind. Coral-reef islands are visible in the 
background. 

Fic. 13. Northern escarpment of Seroe Spelonk (Cl1d), northeastern 
side of Curacao. The foreground illustrates the rather luxuriant vegeta- 
tion near Landhuis Hato; a small, leafless Bursera is conspicuous at the 
left. In the background are the overhanging, older limestone cliffs of 
the flat-topped Hato ridge (Cll, a-d). Although the physiographic 
location of this picture is very similar to that in figure 11, the north- 
facing escarpment is farther from the barren shore and offers consider- 
ably more protection to the vegetation (see location on maps). 


Mo.uuuscs PLATE V 


128 University of Michigan 


PLATE VI 


Fic. 14 (top). South from the summit of the shore-cliffs, Plaja 
Djerimi (C17b), northwestern Curacao. The foreground shows the 
typical, honey-combed exposures of the older limestone; acacias are 
especially prominent at the very edge of the cliffs. The background 
illustrates the very narrow zone of limestone, with the high, undercut 
shore-cliffs, along the western shore of northern Curacao. At the right 
and in the distant background are the higher hills of older rocks, with 
indistinct, fossil beaches on their slopes. 

Fic. 15. Pos Baca (Bel), southern Bonaire. This is one of the 
larger, shallow sink-holes in the recent limestone. Most of the fore- 
ground is bare, due to intermittent flooding, but the background shows 
the same kind of country that is illustrated by the next figure. 

Fic. 16 (bottom). Southern Bonaire (Bl). The brush in the back- 
ground is characteristic of the better protected portions of the low plains 
of recent limestone, south of Kralendijk and on Klein-Bonaire (K1). 
The crotons are especially conspicuous. The foreground is flooded during 
heavy rains. 


VI 


LATE 


P 


cs 


US 


al 


Mor 


130 University of Michigan 


PLATE VII 


Fic. 17 (top). Central, diabasic plains of southeastern Curagao. These 
grassy, rolling plains are practically without molluscan life. 

Fic. 18. To the leeward of Seroe Papaja, southeastern Curacao. This 
is an example of the protected borders of the central plains, where the 
reddish residue in the soil indicates the recent erosion of the limestone 
cap (compare Cb6 and Cb10). A bit of the Hato ridge (C11) is shown 
at the extreme left of the background. 

Fic. 19. Valley between Seroes Palomba and Bientoe (Cb17a), north- 
western Curacao. This is one of the richer valleys of the Christoffel com- 
plex. An epiphyte (Tillandsia) is growing on the brush near the center 
of the picture. It will be noted that much of the vegetation is leafless, 
as is usual during the dry season. 


PuAatTE VII 


Mo.uuscs 


132 University of Michigan 


PLATE VIII 


The scales represent lengths of one millimeter. 

Fic. 20. Stoastomops walkeri. The type specimen from B4. 

Fic. 21. Stoastomops walkeri. Inner view of operculum; the eal- 
careous portion is viewed through the horny plate. 

Fic. 22. Tudora maculata. The type, from B3a, with operculum 
in place. 

Fic. 23. Tudora maculata. Median view of operculum. 

Fic. 24. Tudora maculata. Outer surface of the operculum. 


Mo.iuscs 


SSS 


BRET 
SS 
SSS NUM 


SS = 
2 ae 


G 


ejay!) 
an AD 


A 


134 University of Michigan 


PLATE IX 


The scales represent lengths of 50 microns (.05 mm.). 

Fic. 25. Stoastomops walkeri. Radula from type locality (Bé4). 
Central field and laterals in approximately their relative position; first 
marginal and blades of 3rd and 7th uncini. 

Fic. 26. Cistulops raveni. Radula from C11; except the tilted view 
of the outer lateral which is from C. raveni arubana (A2). Half row, 
with teeth slightly separated laterally. At the right is shown a lateral 
view of the central (R), and median views of the inner (1) and outer 
(2) laterals. 

Fic. 27. Tudora megacheilos. Radula from Cie. Teeth of half row, 
each in position usually seen. Also, lateral views of inner (1) and outer 
(2) laterals, and inner profile of marginal (M). 


PLATE 1X 


Mo.uuuscs 


136 


Fic. 
Fic. 
Fie. 
Fic. 
Fic. 
Fic. 
Fic. 


Fig. 
Fic. 
Fie. 
Fie. 
Fig. 


28. 
29. 
30. 
31. 
32. 
33. 
34. 


35. 
36. 
37. 
38. 
39. 


University of Michigan 


PLATE X 


SCULPTURE OF THE LAST WHORL IN TUDORA 

The scale, shown under fig. 32, indicates a length of one millimeter; 
the rectangles are all approximately one by one and a quarter millimeters. 
Tudora aurantia wassauensis. Female cotype, from B8. 
Tudora rupis rupis. Type, from C2a. 
Tudora rupis newportensis. Female cotype, from Cl. 
Tudora muskusi muskusi. Type, from C17b. 
Tudora muskusi grandiensis. Type, from C14. 
Tudora muskusi bullenensis. Type, from C13. 
Tudora megacheilos, form desculpta. Type, from Cie. The 
sculpture of typical megacheilos consists of similar growth cords, which 
either surmount slightly or extend between the spiral thickenings, which 
are similar to those shown in fig. 36. 
Tudora megacheilos spreitensis. Type, from C8. 
Tudora megacheilos rondeklipensis. Type, from Cl2a. 
Tudora fossor fossor. Female cotype, from Cb17b. 
Tudora fossor arubana. Type, from A2b. 


Tudora pilsbryi. 


Type, from C2a. 


PLATE X 


Mo.uuscs 


Re 


138 University of Michigan 


PLATE XI 


The scales of figs. 40 and 41 represent a length of 50 microns (.05 
mm.), while those of figs. 42 to 47 indicate one millimeter. 

Fic. 40. Tudora maculata. Central and laterals; radula from B4. 
Also outer lateral in profile; lateral view. 

Fic. 41. Tudora muskusi muskusi. Central and laterals; radula 
from C17b. 

Fic. 42. ‘Cistulops raveni. Exterior surface of operculum; from Cdc. 

Fic. 43. Tudora muskusi muskusi. Exterior surface of operculum; 
from C17b. 

Fic. 44. Tudora pilsbryi. Exterior surface of operculum; from C2a. 

Fie. 45. Potamopyrgus parvulus. Large, globose form, from Bel. 

Fie. 46. Potamopyrgus parvulus. Elongate form, from Kel. 

Fic. 47. Potamopyrgus parvulus. Spinulose form, from Kel. 


Mo.uuscs PLATE XI 


140 University of Michigan 


PLATE XII 


The hair-line indicates a length of one centimeter; the dimensions of 
the shells figured are given in the text. 

Fie. A. Tudora aurantia aurantia. Male and female, from B2. 

Fic. B. Tudora aurantia wassauensis. Male and 2 females, from 
B8; the right-hand figure is the type. 

Fig. C. Tudora muskusi muskusi. Male and 3 females, from C17b; 
the second figure from the left is the type. 

Fic. D. Tudora rupis newportensis. Male (type) and 2 females, 
from Cl. 

Fie. E. Tudora rupis rupis. Male and female (type), from C2a. 

Fie. F. Tudora muskusi grandiensis. Two males and two females, 
from C14; the right-hand figure is the type. 

Fic. G. Tudora muskusi bullenensis. Male and female (type), 
from C13. 

Fie. H. Tudora muskusi X Tudora fossor djerimensis. This speci- 
men appears to be a hybrid between these two species. The operculum 
is also somewhat intermediate, but is much closer to muskusi. From C17b. 

Fic. I. Tudora fossor djerimensis. Male and female (type), 
from C17b. 


Mo..Luscs PLATE _XIT 


142 University of Michigan 


PLATE XIII 


The hair-line represents a length of one centimeter; the dimensions of 
the shells figured are given in the text. 

Fic. A. Tudora megacheilos megacheilos. Male and female, from Cde. 

Fic. B. Tudora megacheilos, form desculpta. Male and female 
(type), from Cde. 

Fic. C. Tudora megacheilos spreitensis. Male and female (type), 
from C8. 

Fic. D. Tudora megacheilos rondeklipensis. Male and female (type), 
from C12a. The male is turned so as to show best the palatal emargina- 
tion of the peristome in T. megacheilos. 

Fic. E. YTudora megacheilos kabrietensis. Male (type) and female, 
from C3. 

Fic. F.. Tudora pilsbryi.. Male and female (type), from C2a. The 
male is turned so as to show best the palatal sinus of the peristome in 
this species. 

Fic. G. Tudora fossor fossor. Male (type) and female, from Cb17b. 
The type retains all of the whorls. 

Fic. H. Tudora fossor westpuntensis. Male (type) and female, 
from C18. 

Fic. I. Yudora fossor canashitensis. Male (type) and female, from 
A4b. The dimensions given in the text were taken when the type retained 
all of the whorls, but a bit of the apex crumbled away before the figure 
was made. 

Fic. K. Tudora fossor arubana. Two males and a female (type), 
from A2b. One male (the left-hand figure) has the spiral sculpture very 
much reduced; the other is turned so as to show best the slight palatal 
emargination of the peristome, which is usually greater in arubana than 
in the other subspecies of T. fossor. 


PLATE XIII 


Mo.Luuscs 


144 University of Michigan 


PLATE XIV 


The scales represent lengths of 50 microns (.05 mm.). The centrals 
and lst laterals are shown in their proper relation to each other; the 
other teeth are simply oriented in regard to the long axis of the radula. 

Fig. 48. Suecinea gyrata. Radula of a medium-sized specimen from 
Cde. Central, 1st, 7th, 14th and 21st teeth. The three examples of the 
14th tooth oceur within four, consecutive, transverse rows. 

Fic. 49. Succinea gyrata. Jaw from same specimen. 

Fig. 50. Thysanophora vanattai. Radula from A38. Central, 1st, 7th 
and 14th teeth. 

Fic. 51. Thysanophora vanattai. Jaw from same specimen. 

Fic. 52. Drymaeus virgulatus. Radula from Clle. Central, both Ist, 
3rd, 28th, 52nd, 54th, 57th and 84th teeth. The hair-line below the scale 
shows the shape of a transverse row. 

Fic. 53. Neosubulina harterti. Radula from B5. Central, 1st, 7th, 
14th and 19th teeth. The shapes of the smaller cusps of the outer mar- 
ginal are, of course, indeterminate, as their diameter is less than half the 
length of light-waves, so they are only visible as points of light. 


146 University of Michigan 


PLATE XV 


Each scale indicates a length of one millimeter, except in fig. 55, where 
it represents 200 microns (.2 mm.). 

Fic. 54. Guppya molengraafi. Apical, basal and profile views of the 
type, from Cb20. 

Fic. 55. Thysanophora crinita arubana. Profile view of hair and de- 
tail of sculpture on 3rd whorl of a young shell from A4a. 

Fic. 56. Thysanophora crinita arubana. Apical, basal and profile 
views of the type, from A4a. 

Fic. 57. Thysanophora vanattai. Apical, basal and profile views of 
the type, from A4a. 


148 


University of Michigan 


PLATE XVI 


The scales in figs. 58-63 indicate lengths of one millimeter; those of 
64-66, 50 microns (.05 mm.). 


Fig. 58. 
adult (B6). 
Fic. 
Fie. 
Fig. 
Fic. 
Fic. 
Fic. 
Fic. 


59. 
60. 
61. 
62. 
63. 
64. 
65. 


Neosubulina harterti. Embryonic shell, from body of 


Neosubulina gloynii. Ditto (C17). 

Neosubulina scopulorum. Ditto (A3). 

Neosubulina harterti. Largest adult (B6). 

Neosubulina gloynii. Largest adult (C11). 

Neosubulina scopulorwm. Type, largest adult (A3). 
Neosubulina harterti. Jaw (B6). 

Brachypodella raveni raveni. Radula (C5); entire view of 


central and tips of two others (C), inner lateral (1), outer lateral (2), 
1st and 3rd marginals (3, 5). The teeth are simply oriented in regard 
to the long axis of the radula, and are not shown in relation to each other 
(cf. Pilsbry, op. cit.). 

Fic. 66. Brachypodella gibbonsi. Radula (B3); tip of central. The 
scale also applies to the preceding figure. 


PLATE XVI 


MoLuLuscs 


150 


University of Michigan 


PLATE XVII 


The scales indicate lengths of one millimeter; figs. 67-71 and 74-77 
are drawn to the same scale. 


Fic. 
Fic. 
Fic. 
Fic. 
Fig. 
Fia. 
Fic. 
Fic. 
Fie. 
Fic. 
Fic. 


67. 
68. 
69. 
70. 
Tale 
72. 
73. 
74. 
75. 
76. 
Utle 


Brachypodella raveni raveni. Complete adult, from C11. 
Brachypodella raven sanctaebarbarae. Type, from C2a. 
Brachypodella raveni knipensis. Type, from C17a. 
Brachypodella raveni arubana. Type, from A2b. 
Brachypodella gibbonsi. Type, from B3. 

Brachypodella raveni knipensis. Conical specimen, from C17. 
Brachypodella raveni raveni. Apical whorls, from Cdc. 
Microceramus bonairensis bonairensis. Topotype, from B2. 
Microceramus bonairensis curacoanus. Type, from Cde. 
Nicroceramus ‘bonairensis curacoanus. Specimen from C17. 
Microceramus bonairensis arubanus. Type, from A4b. 


XVII 


2 University of Michigan 


PLATE XVIII 


Variation in Cerion wa uva (Tables XI, XIT) 
Fies. C6, D5. C. uva desculptwm (Cb6, Cb10). 
Fic. A2. Type of form diablensis (C12b). 
Fic. F6. Type of form hatoensis (C1l1d). 


Mo.uuscs PLATE XVIII 


a | 


Verne ON any 


: 


eee 


” 


Anal 5 we pili 


bz 


i ASS ry 
\ eueet| 
Mays) y 
eee 
‘Teta | 
wat. 


EO aes 


154 University of Michigan 


PLATE XTX 


Variation in Cerion uva kiipensis 

Fic. Al. Type of form djerimensis (Clic). 

Fics. C3, 6, D3, 4, 6, E4-6, F3-5 from substation Cb17. 
type of knipensis. 

Fic. C5 from station C20. 

Fies. D2, E1, 2, F2 from station C18. 

Fic. E3 from station Cb16. 

Fic. F6 from station Cb20. 


F5 is the 


PLATE XIX 


Mo.Luuscs 


156 University of Michigan 


PLATE XX 


Variation in Cerion wea arubanum 


Scale and arrangement as in Plate XVIII. For extremes, 
Table XIV. 

Fics. Al-3, B1-3, C1-5, D1-4, F2, 3, 6 from the colony at A2c. 

Fic. C3 is the type of the subspecies. 

Fics. A4, B4, 5, D5, 6, E4, 5, F3-5 from colony at Ada. 


see 


Mo.uuuscs PLATE XX 


158 University of Michigan 


PLATE XXI 


Variation in Cerion uva bonairensis 

Seale and arrangement as in Plate XVIII. For extremes, see 
Table XIV. 

Figs. Al, B2, 3 from Station B9. 

Fic. A2, the type of form kralendijk, from station B1. 

Figs. A3, B1 from station B2. 

Fies. A4, C2, D3, E3 from station B4. 

Fics. B4, C5 from station B7. 

Fic. B5 from station B8. 

Fies. C1, 3, D2, F2 from substation B3a. 

Fies. C4, D5, 6 from station Bb7. 

Fies. D1, 4 from Klein-Bonaire (K1)._ 

Fics. E2, 4-6, F3-6 from station B5. F6 is the type of bonairensis. 


PLATE XXI 
Fr 
\ 
\ 
WANN 
\\} 
7 | 
~, 
‘ \Y 
\ 
™>, 


> 


INDEX 


PAPERS 129—152 


New names and numbers of papers are printed in heavy type 


A 

Ta SLTETES ete Re Ce le 152, 10 
ORB NU OSS eet See 152° 10 
JOD ATE a ee 130, 32 
Adamsiella aripensis................ 152, 42 
Adelopoma occidentale..137, 46, 47 
[MECAET OS, See ie ty eee 137, 48 
LETS i cre 151, 2, 12 
gL el ee 152, 11 

Agnurodesmus thrixophot.............. 
sh, va Reshma 133, 3, 82, 83 
MBETHCOSUNS: 42 tt... 133, 83 
Joo) 0 ee 134, 1 
FPS ATID AM ER een 134, 1, 7 
AasSSOCGESMUS. .-rec.-sssesssessssessteteseee: 133, 59 
PeeNGtES 133, 2, 59 
ESNNALED ESR oS le ee a! 137, 13, 14, 15 
Alsophis cinereus... 132, 6 
IGECSMAOl ee. 132, 7 
TIVE Nak oe ee 132, 7 
Ambloplites rupestris................ 138, 2 
PEEVE BIER (ooo 132, 1 
AGT Ci 1G) Ct: rr 138, 2 
TASC 138, 2 
TCM 423k ee 138, 3 
LLIN i 149, 1, 3 
bifrontata ........ 143, 7; 149, 1, 2 
GEST) ae ee 143, 8 
erythrocephala .....:cccccc- 132, 6 
E10) 0 OE ae 132, 5, 6 
FLT AUT Ch SN a 129, 9 
2 US ee ae 132, 6 
LDS CLG De ase eae 132, 6 
MEPSTPEATI CA oe nie 149, 1 
ARG VISATRA fe orcas 132, 5, 6 
Quad rilimeata nn eeccsecssnes 129, 9 
EER CF EEL PUES nesses, cee hs 129, 9 


SAGER sCALV eon een 138, 2 
Amphicyclotus 
ge he 137, 28, 29; 152, 112 
boueardi 
cayennensis 
lutescens 
ponderosus 
Annularia lachneri 
aurantiaca 
Anolis 
aeneus 
alligator 
barbudensis 
bimaculatus 
binotatus 
bonairensis ..148, 4, 7; 152, 29 
cepedii 
forresti 
gaigei 
gingivinus 
homolechis 
latifrons 
limifrons 
lineatus 
princeps 
roquet 
squamulatus 
BIDOMIOSUS hee te .129, 
wattsi 
Antirrhoea acutata............. 152, 10 
Aperostoma bartletti................ 137, 32 
confusum 
connivens 
sanctaemarthae 
smithi 

Aphelidesmus guianensis............. 
cae ERS ah ee 133, 3, 46 


or we: i) ee | 


SANA AaNEAKrE H 


mm D 


2 University of Michigan 


Aphredoderus cence 138, 1,3 
Say anUs 8 138, 1 
Archaeogomphus.....134, 1, 5, 6, 7, 8 
hamatus..134, 1, 2, 3, 4, 5,7, 8,9 
furcatus 
ERAN SS Sot oe eee 134, 7,8 
Argyrosomus prognathus......... 146, 1 
Pia tetigy et ee 133, 75, 77, 78 
ulophilus ........ 133, 3, 75, 76, 78 

UM 45) () Saas seat i ee eet 150, 1 
elegans 
elegans___— 150, 1, 2, 3 
occidentalis... 150, 1, 2,3 
PERCU 6 129, 11 


ee Sees A 29518 AS 
certs es 129, 12, 13 
WARIHS) oo ae. 129, 11, 12, 13 


Batodesmus ................ 
Boleosoma nigrum 
Brachypodella ..152, 91,93, 112, 114 
gibbonsi .......... 152, 94, 111, 114 
hanleyanen. 2 2 ee 137, 26, 27 
raveni_137, 7; 152, 90, 111, 114 


Pave oe a ee 
es 152, 90, 91, 92, 93, 94, “4 
sanctaebarbarae —— 
Sse enee tee 152, 91, 94, 113 
Branchinecta occidentalis...141, 1 
Brachypodella raveni................ 137, 7 
Bromelia lasiantha................. 
PSNORNG YES cos ee 
Bufo haematiticus............... 
MAPMNUS °c 129, 10; 132, 7 
sternosignatus 
typhonius 
Bulimulus apiculatus.......152, 80, 81 
COTIBCBUS on. 135, 9, 10 
dlongatus  _ 152, 80 
Bulimus elongatus..................... 152, 80 


Multilineatus een 152, 108 
torallyi sisalensis........... 152, 108 
Bumelia obovata.n ccc 152-08 
Bursera 22 ees 152, 11, 24 
gummtfera, 152, 80 
SCNNTHDA 2 152, 11 
Cc 
Caesalpinia coriaria................... 152, 10 
Casearia bonairensis 152, 11 
Calladinm, =. 133, 9, 31, 35, 47 
Capparis Breynia.................. 152, 10 
jamaicensis 
Catostomus commersonii.......... 138, 2 
Cereus, 2. 152, 10, 16, 17, 61 
Cerion 40 eee 
137, 2; 152, 13, 16, 18, 109, 
111, 112 
SNICOUIL) = 152, 111, 112 
APIAPOM a2 152, 98 
Wy! 
137, 7; 152, 36, 98, 99, 
104, 110, 115, 116 
atubannim. -. 
152, 98, 104, 105, 106, 114 
bonairensis,) ———___— 3. 
152, 98, 105, 106, 114 
desculptum, —..=__ 
137, 7; 152, 18, 98, 100, 
105, 106 
knipensis —____ 
152, 20, 98, 102, 103, 104, 
106, 114 
kralendijki 152, 98 
uva 
152, 18, 23, 27, 98, 100, 
101, 102, 103, 104, 114 
vulgare (2. ee 152, 98 
Chaenobryttus gulosus.............138, 2 
Gliara 2 eee 138, 2, 4 
Characodon atripinnis................ 148, 2 
PLLC A TUS, cee 148, 7 
ferrugineus <== 148, 3 


Occasional Papers of the Museum of Zoology 3 


Anvepoldty See eee 148, 2 
Wariabus) ct en 148, 3, 4 
Chondrodesmus armatus.......133, 69 
CETASINOPUS...----eeeceoe 133, 3, 69 
eth 4)2 4 a8 1: eee neat 133, 69, 72 
EUSOSIO’ <5 ots 133,73, 71 
tamocalanus........... 133, 3, 67, 72 
virgatas—......133, 3,70, 71 
frater:.. 133, 3, 71 
Phondropama 137, 24 
PAVGHN = ose: 137, 4; 152, 37 
SUDAUTICULALOIA aosesccsssessssennsesenssee 
LRG eee eee 1375-2605 152) 112 
Cionella gloynii.......... 137, 5; 152, 88 
Cistula maculata 152, 116 
TET TLE Oe eae ee 152, 37 
Cistulops.....152, 35, 109, 111, 112,115 
TED YC ee eee ee 152, 113, 116 


arubana...152, 36, 39, 40, 114 
raveni..152, 36, 37, 39, 40,114 


BH Gd ests oe ste iane 133, 79 
cryptopygus........... 133, 3, 79, 80 
Cnemidophorus arubensis.......143, 8 
LES Tadeo 710 (7 tes eee one 143, 8 
STE ee ee 143, 8 
VOMMNISCATUS oecennccsessscssee 143, 8 

TPP TACIT (2 DA ve ree 143, 8 
PETAR OLOE iene tenses seees 143, 8 
Coccoloba uvifera...................... 152, 23 
Coecilia sabogae....................... 129, 14 
WOIGDOSL YUE oii. ceecsntccscsten. 152, 41, 42 
Colombodesmus.............. 133, 53, 56 
etd st-hg) 133, 2, 53, 56 
DRT Sirtcee tease scstnisase anni 133, 2, 55 
“OLE ear ee ee ee 1395, 1,12 
constrictor constrictor.....139, 2 
AVEVONGFIS 26.580 fise tessa 139, 2 
STEC 1 tech; s Bepeepeeee eo 139, 2 
(DUCT U1 eee ee ee 140, 1 
CDEC peal ec pence OOS or 139, 2 

ME GETTIO CLE SETS nse ssesscnsesssssnccsecoveees 133, 56 
[Tob arse iol fb toe eee 133, 2, 57 
Corythophanes cristatus........... 129, 4 
(4, 711 Sia eer ee a a ee 130, 45 


acutus 
floridanus 
lucius 

Crotalus horridus unicolor.....143, 9 
terrificus ... 

Croton) =.= 
flavens .... 

Gy clanthtis. 726s. 130, 6, 22, 
biparhitua: 130, 23, 25 

Cylindrella raveni................. 152, 90, 94 

Cycloporiare se eee ees, 

Cyclostoma aspratilis 
aurantiacum................. 
DEEQRING esse ction: 
blanchetianum.............. 
POMOUGHSISi Sten eee ee 
PEASUHENSE), 25. .en re 137, 28 
eancellatum................... 152, 56, 58 
CANUGIIN ten 152, 45 ,46 
CUE UA TOV oe oenceene onan 137, 42 
COLOMDIENSAS «..--ceeccssseseeceeteeee 137, 44 
CUMANEDSE.....eaaennse—---- 137, 23, 26 
CUIMITED TE eee cee eso, 137, 42 
GiSCOTd CHIN Ieee. Scien 137, 29 
disney 2 ee 137, 29 
GistinGhum We. ete 137, 29 
PIP ANEW Fee cath ee 137, 42 
glaucostomum ow... 137, 37 
@TANACENSE vies 137, 37 
TNCs eee eee 137, 30, 44 
INCOM PLUM, 2 eee 137, 30 
IMCONSPICUUM 0... 137, 44 
TPT OP UAT Oe acters 137, 41 
Nepean cece 137, 41 
mepacheilos) [asco 152, 55 
mepachevlomiy ncn. 152, 55 
megacheilus .......n..-- 152, 55 
NO DVO pec eee tee 137, 44 
PlAMOT DUO ccc 137, 28 
plicatulum.................. 137, 23, 25 
POP RY A ee ce ssccsssoese 137, 44 
POF OMNEN TE ac cssssccctesen es 137, 28 
DP OUGUS Ret dectancn cee csmcs 152, 56 


University of Michigan 


psilomitum.................... 137, 28, 29 
({UELENIBES Soe eqn chee ee 137, 43 
TILA WA dco tentna cena 152, 42 
FOLIC 2248, 152, 55 
PIM Ces te che a 152, 41, 55, 58 
stramineum.................. 137, 30, 37 
SUPHEALG eee 137, 29 
PANGAN eee 137, 23, 25 
translucidum.................. 137, 29, 33 
fgg en Retelsth eens per eee 137, 33 
VETLEZUCLENSE......ccececessseee 137, 23, 26 
WVETSICOLOT 2 jens 152, 45, 46 
Cyclotus berendti 0.0... IST Ido 
bisinuatus 32 
boucardi 29 
COO UGE oa caer cee 29 
COPPUWLEWGUS «sciences ate. 137, 43 
unikerit stn ne 137, 43 

GY SOW eee racentreactice 137, 34 
AUDI Geen eee 137, 34 
AMDIOUUS! (iat acess 137, 34 
TWIN OF ott ence eee, 137, 33 
multilineatus.................. 137, 33 
floltraAtus 2a... 137, 41 
ASCHOTIS Serica ono 137, 41, 42 
SPANMIATUS! on oceeressin 137, 37 
irregularis pittieri......... 137, 41 
DANG seer tei cts ena 137, 41 
POEOTN hage cibnehieieciee: 137, 43 

TUG AGUS A pecietuciese Cua 137, 37 
Cyr OdeCSMUsi ce res 133, 79 

D 

Decodon verticillata.........0. 138, 2 
Dendrobates lugubris................ 151, 5 
TATICLOVANIS conan or 129, 13 
DTA O PIB 32 ccreseecrienartregnctiaroncen 142.1, 7 
PUNYE AU ULI eerie eee 142, 3, 7 
ehccks| opi it ey neem 142, 3, 4, 8 

TOC OSUNS! uhacen ance 142, 8 
occidentalis............----- 142, 6, 8 
PUICHENUS......eeceerereene 142, 6, 8 
BRISTLE NG iescstcstssersesisccaeseonoe 142, 4, 8 
vandenburgii......142, 5, 6, 8 

POO ANAS eisai itrntimseroert 142, 1, 2, 7 


arizonuae.. 2 ae 142, 2,7 
TOG alig eee 142, 1, 2, 7 
Diaphoranolisi<-o- see 129, 7 
brooksi... 129, 7 
Diploglossus monotropiz........... 129, 3 
resplendens 2275 129, 4 
IDOcoOdeRmINSy .25--e eee ee 133, 75 
Dromicus antillensis .0000.. 143, 9 
Dromodesmus. »rreciccoscssssvserseee 133, 5L 
LON GIPES...reeeeseeesseee 133, 2, 51, 52 
Drosophila’ 225.000). . eee 132, 8 
Drymaeds’ 2.55 ee 135, 1 
DYYMeu....ceccccsecsesene 135, 1; 152, 111 
AlbostTiatus......creeee 135, 11, 12 
COMINICUS....... eee 135, 10, 11, 12 
INGOT PUNCEUS 2 eeceeconses 152, 85 
multilineatus =. . ee 
135, 12; 137, 6; 152, 108 
virgalatus <2)... cee 


152, 14, 21,22, 80, Bape 
84, 109, 110, 115 


E 

Eleutherodactylus palmatus.......... 
Sedo tntyccde te te ea 129, 10 
TANOIMCS) = ..cee eee 129, 10 

Elliptio plexus crocodilarum........ 
Sc cierto ee 135, 1 
crocodilorum .............. 135, 1 
Hlodea* acl ee 138, 2 
Hpinannolene 2p 133, 19 
ALIWG ice ee 133, 2, 18 
LOLENZONUS -:rreeeccssseeeeeeeeeee- 133, 2, 16 
Testis 3 133, 2, 17 

Epistreptus eustriatus.--.cccccccc0 
Bias Mine basen bee 133, 2, 21, 26 
FOYT YZOT SUCCOEB uereccsenesersssserssecns 138, 2 
Erythrolamprus aesculapii..129, 14 
Esox vermiculatug............00. 138, 2, 3 
Eucalia inconstanag............. 138, 1, 2, 3 
Mn comulis 2 ec cennceaee 135, 15 
elegantula .... 135, 1 
C-) (4211121 1: a 135, 1 
Euglandina decussata....ccn 135, 8 
PONG dois oactauniioea yee 135, 8 


Occasional Papers of the Museum of Zoology 5 


Eupemphix pustulosus.............. 129, 10 
Eupomotis gibbosus.......... 138, 2, 3, 4 
ENT EROCHATCN A Scot osccnntantees 152, 34 
SOMA hs rece eno 152, 34 
PLUMP OTIANA a... ceosseetceeenesaen 152, 35 
semilirata........1387, 23; 152, 112 
Evermannichthys.............00:0 144, 1, 2 
metzelaari --........... 144, 2 
BONO UCOl a ese ceerctcinnsn 144, 1, 2 
F 

Ferrissia michiganensis ...145, 2 
TUES Wi ace eee ona 145, 3 

ri) 2ietG C7 A Re cae ee al ee 145, 3 
[DICCSTIS) De Se a Re een 152, 21 
Fundulus notatus 2... 138, 2, 3 

G 

UAE ee eee br 144, 1 
Bponmcols 144, 1, 2 
GAStTOCOPEA e--seseernencseeee 152, 74, 116 
barbadensis hojeda .....152, 117 
BTA O AT ences 152, 116 
longurio ............ 152, 74, 116 

IG OYE a reece 152, 117 
Glomeridesmus orphnius ......-...-- 
EE Oe 133, 3, 4 
moreellus: 133, 2, 3, 4 
Gobius ParadOxXus -2...-ccccecceecee 144, 1 
Gonatodes albogularis ............. 143, 2 
fuscus 2 
vittatus 2 
RRO LC ce sence 7 
atripinnis ....148, 1, 2,3, 4,5, 6 
PORES) ooo ee 148, 8 
CORLISS ale oer ee ae 148, 3 
SOTA aie ete. 148, 4, 5 
MET U Dee e h ee 148, 8 
tuitpoldi 148, 2, 3, 4, 6 
Ly ee en ee 148, 4 
EsEUZIEEY OTE Ce), oie ne ae ee 152, 24 
WLU NS iss ccancarst 152, 11 
ReRPETIOTUS ns 133, 77 
Sctoporus..3. 135,) 3, 17 


Guppya ..____ 135, 15; 152, 111 
gondiaeht 2202S. 152, 77 
MIGMUONSIS!: 2 152, 77 
molengraaffi..152, 76, 110, 114 

Gymnodactylus antillensis .....143, 2 

Gymnophthalmus quadrilineatus 

eee ee eee 143, 9 

gts is LF gestae oe Rae ene hak 152, 74 

H 

Prabrocontis) 22s oe 135, 15 

Helicina braziliensis ................. 137, 18 
GANGGAN ain ee eee 
earacolla, .......... 
christophori .... 
colombiae ......... 
columbiana 
concentrica 

concentrica. ............ 137, 16, 17 

pandiensis ............. S74) 16, 17 
Crassilabris 2. eer 137, 21 
TOnOCHe ane ees 137, 18 
gonochila 22 137, 18 
ADR S'S] AER Sees eed ero toon Ps 137, 18 
kieneri —....... 137, 21; 152, 112 
MOMOLAL Sees eee 137, 21 
OCANBHEIN ee 137, 18 
THOGOSCOMA ae seesseeesseneteee 137, 21 
ThynchostOma, -....cccceeceeeeen 137, 18 

ernesti 

infesta 

rhynchostoma, .............. 137, 18 
TID SEA ere 137, 13 
sanctaemarthae ................. 137, 20 
steindachneri ............ 137, 18 
SUPOLSCLUCEA, anne neocon 137, 18 
neil cv P12 h s oators eee neea cee eee 

yA eee dey, toy a7, 19,20, 21 

BU PUT ee eos 137, 19, 20 

TANIA Se eon 137, 20 
AO) OF: ht: Rene, steel eee 137, 21 

Helix elongata 2... 152, 80 
incerta, —..... 152, 107 

MLO LUD UNIS pee scene 152, 107 


Bg a ae 152, 107 


6 University of Michigan 


notabilior _______152, 107 
notabilis ________152, 107 


Indoviea ______152, 80, 81 
virgulata _______152, 80 
Hetaerina ____130, 1, 6, 7, 8, 9, 
10, 11, 12, 18, 21, 26, 29, 30, 

32, 34, 39, 41 


americana 130, 12, 13, 
20, 21, 33 
caja ____130, 8, 9, 10, 12, 


13, 15, 16, 17, 19, 21, 22, 23, 
24, 25, 26, 27, 29, 30, 31, 32, 
34, 35, 36, 37, 38, 39, 40, 41, 
42, 43, 45 
eapitalis 130, 8, 9, 10, 12, 
14, 15, 23, 24, 25, 27, 29, 31, 
35, 36, 37, 40, 41, 42, 44, 45 
eharea__130, 12, 14, 19, 28, 29 
cruentata._.130, 12, 14, 15, 17, 
21, 27,31, 42 
dominula 130, 12, 15, 19,39, 46 
fuscoguttata___.130, 12, 16, 39 
laesa___130, 11, 12, 16, 39, 46 
 * ees Was 
maeropus 130, 8,9, 10, 12,13, 
14, 15, 16, 17, 18, 19, 21, 23, 
24, 25, 26, 27, 28, 29, 30, 31, 
33, 34, 35, 36, 37, 38, 39, 40, 
41, 42, 43, 44, 45 
miniata 130, 9, 10, 12, 15, 
18, 29, 32, 34, 36, 38, 39, 42, 
43, 44, 45 
cel! i a 2 
130, 11, 12, 18, 19, 39, 46 
mortua.__ 130, 11, 12, 19, 46 
pilula._....__130, 12, 19, 43 
sanguinea —___ 
ae ae 130, 12, 19, 28, 29, 42 
titia__130, 12, 13, 17, 20, 33, 
34, 36, 38, 43 


meerelage 85 2 ee ee 
Heteragrion «130, 10 
Caryn 4 de, 
it | * | enemies: Fal | 


Himantodes cenchoa 129, 14 

elegans __ +. see 
Hippomane mancinella_._.__ 

eunennmnenee: e 

Hyl, ee 

albomarginata.__..__151, 2, 3 


baudimii  ________ 43 
dolomedes —————____129, 11 
cahia eee 
Mmaxma eee 
uranochroa ___ 1 ae 
I 
es: ree 
iguana __._____.__ ie 
delieatissima _______132, 5 
ieuana ee 
Impatiens bifiora 138, 2 
Tris versieolor _._______ 138, 3 
J 
Jatropha urens________152, 25 
L 
Labidesthes siceulus__.____138, 2 


Lantana 1 
Lathrogecko sanctae-martae. 
aoe eee 
Leimadophis trisealis_____143, 9 
Lepomis ineisor____________138, 2 
Leposoma —______._ #7 a 
dispar —__._ eee 
southi—___..__14732 ee 
taeniata 147, 1 
Leptinaria gloynii__137, 5; 152, 88 
minuseula 137, 5; 152, 88 


harterh 
helenae __________ 158, 8&6 
martensi —___._-_ ae 
Mexieanum —.__._____135, 8 
triptyx ____—____12, 86 


Leptoealamus torquatus.__129, 14 
Leptodactylus bolivianus__129, 10 


Occasional Papers of the Museum of Zoology if 


TOUT UMEH AVEO Ces mmeetenerar 
melanonotus .......... 
pentadactylus 
Leptodeira anuulata.... ccc 
Chea Speen 129, 13; 143, 9 
QU CICHIGDIYS) s..::fececccrvsrtetcenses 146, 1, 5, 8 
alpendeacncercasit 146, 1, 5, 8 
UELOGM wercanaerertemer catia nett 146, 6 
VG} 2b rector ee ca gereteromeesoen ite 146, 5 
IGG eee te een, oe 146, 5 
SJ OMAMMAG sesh ine tecensicts 146, 1 
1 enifed 0 ga 6 reereeeeer reese 146, 5, 7 
1 Uist] fe V Gib), eect aetna eee 130, 12 
. quadrimaculata. ............... 130, 3, 4 
TNO SIS|AD sberertht bare baa 130, 4 
TLSVOFESS SS ee bs ieee ae a 152, 42 
Liguus virgineus ..........0c.- 152, 108 
DFC CATS AY WE eee ieee 137, 22 
POUT COND) Ketter teeter 137, 23 
IIL TU, epee eee che Re 137, 22 
aM ll O Saree cet csiatiees 137, 22 
Lr 2,702 sp eae ae 137, 22 
plicatula christophori................ 
_rtmenc tLe eect eee ERE ORR 137, 22 
venezuelensis «0.0.00 137, 21 
Lymnea columella ou. 145, 3 
humilis modicella................. 145, 3 
M 
Machaonia ottonis. ........ 152, 11 
Macroceramus inermig.............. 152, 96 
UTP CEN ATAU rscsastssssctessesnctinscccncsanesanece 138, 3 
INE SHE Glo) OL 0a a are eo er 139, 1, 2 
EUTIMAINO TRV Mee gee ctarcecitntectesncearazess 139, 2 
PANUTU OUTS ectsk te cectts net stetectet 139, 2 
TORONTO ccs earner: 139, 2 
flagellum flagellum............. 139, 2 
LiLT haben EW to) Dyanna ee 139, 2 
PSE TUPUUTIS inarso ds Seas voncesssssrs 139, 2 
URIS EULESS meee trrsecrctecsenstrisar reser 139, 2 
RIMS A eee nad ete ianie 139, 2 
GMO VANIUA vecratacctveett 139, 2 
PICOUS sete ee cstanncasrneventaaes 139, 2 
ruthveni ..... .139, 2, 3 


SOMO tLe ee Us} heresy ry 1 
SCMIUINCACUS oanseeeecccsressseecereeeen 139, 2 
GACTTAG UR! si. sctoesncasteceee areca 139, 5 
PAP ANN cpa cre tee naccteorerne 139, 2 
GELCTUVAUUIS ercecrer este esses tsassec te 139, 2 
Me malo HI Se essere ctascecmsseesteens 151, 12 
AMONPAN A: vata ccmernnd Asn akan, i 
MiclOCHeTUS!  arics.c0- nee 152, 10, 11 
Melo Chi areas eee eee 152, 10 
MiCroOCeEraMUs oeeeerecssccceeenn i Ns y2eea ba Lak 
IDONAITENSIS Rn. caetenacteser: USW(- eff 
arubanus........... 152, 97, 114 

IDOWEATOTISIBG cecpeete cere cscs 
eee 152, 95, 96, 97, 98, 114 

CUT ACO AT Are iocctoiccstess 
bate aE 13%) Osi se tos, 96 

CHITA COAMMS ieee ets: 
ees 152, 96, 97, 98, 114 
pontificus............ Ti towels, OF 
Microhyla achatina ..........0.. ilaahs. 7/ 
Microperea punctulata............. 138, 2 
Micropterus salmoideg........... 138, 2, 3 
Microspirobolus tridens...133, 2, 41 
MT CriIPUsee a eet carci 129, 14 
CUSHOLGUCUS Meenas ecreens 129, 15 
CUTER ee ernetaae ene 129, 15 
WO etnies ve ccrense ten 129, 16 
TFT OCW CUTS Mee. ned 129, 14 
Tif S(OL CWI YG i praetor ic aera Sees er eee 129, 15 
Mimulus jamesii .........c cen 138, 2 

N 
Nanostreptus astix.....1383, 3, 21, 23 
PTAC IOe eyes 133, 21, 25 
incertelineatus 


INCONStANDS .o........... 


confluens........... 
fasciata............ 
pictiventris............. 140, 4, 5, 
ING OGY CLOUIS! rcs csetcccececcseene 137, 28, 29 


8 University of Michigan 


DOM state ann cect antares 137, 42 
CANICAENSISE nan nana 137, 41 
CHTYVSACINE) ei terraces 137, 29 
COLOMpIENSIS sae. 137, 43 
GEPLGSSUS Hs neta 137, 42 

giganteus subcingulatus....... 
See ann he ee eae 137, 42 
AMAL AGUS seem eere es 137, 29 
PANAINENSIS ss sees 137, 37 
POW SL: visi. ch caswnramamanes 37 
pergrandis 30 
peruvianus 42 
popayana fasciata........... 137, 44 
SOlMGIS. eee ene eee 137, 29 
Neosubulina.....152, 85, 86, 109, 111 
gloynii........... 152, 87, 88, 89, 114 

har Ger fileen er canaces aaa art 
137, 5; 152, 27, 85, 86, 87, 

88, 89, 114 

SCOPUlOLUIN |e 
eee 152, 87, 89, 90, 114 
NOT aie Gotan 152, 33 
Pesselllataqvnecea enon 137, 13 

Neritina virginea microstoma..... 
mente treater Rane teasereae 137, 12 
MEClivaibae nacre 137, 12 
GIO oattecetan ainenao tse 152, 32 
Nocomis kentuckiensis.............. 138, 2 
INOTOWSi ea caae enantio natesetst 129, 7 

Notemigonus erysoleucus.........00« 
eth d che Mae enn 138,°2; 3 
Notropis Cornutus...........--0 138, 2, 3 

O 

Wve ci natea oer 137, 14, 15 
CACALUCIITAevacdccseete: 137, 15 
dysoni barbatia................. 137, 15 
CV SOUT Rita strate ee 137, 14 
POU 5. ssilacucnegorutnen 137, 15 
PUP ATLA Aa ncnaesae cme 137, 13, 14 
tachirensis........... 137, 13, 14 
MOLTO ataaetaemenunetnn 137, 15 
TELE MEN crececaciasiinacaTntante 137, 15 
MONICOAa za tere tater 137, 13, 14 


Lq(b(e) ey Bere cepctaree nee carecrete 137, 13 
PATRCNSIS: ‘ec. scoacat 137, 13 
OPGASiFesceosscstastesecn reer 152, 86, 87 
beckianum........... 135, 8; 152, 87 
PTAC Ecc teaicrornne nee 135, 8 
PT CT A: 2.:cdnaisnakteentae 152, 108 
Opuntia ees 152, 10, 21, 25, 26 
Orthoporus, etholax:.........c0s.ne0u 
ies 133, 3, 21, 30, 31, 32 
foliatus eee 133, 3, 21, 34 
gaigei 133, 2, 21, 32 
FS] OSes er earn eraeeere ara! 133; 3530 
walkeri 133, 3, 21 site 
Otala aspertlagenerccra 152, 107 
(0>.9'4:119'd Cree rneereer 
crossel ..... 
ferussaci .... 
TONGA), Beis ca daiacanenas 
maracaibensis 
imitator 
PUINCEPS vacate 135, 12, 14, 15 
EYICI GEA) eeateeeecestacnd 135, 13, 14 
ETI ASCLAL AN cee oneee 135, 13 
PIE ACtA cece eass ern 135, 13 
Pp 
Paludestrina crystallina........152, 70 
Valenciae cute naeatey 
Paludina coronatus 
JAMAICENSLS..........-nen 
DAT VW) crctunneanthes 
Pantala hymenaea 
POrip ats), vaitcccd.cstenanomemneen 
Perithemis | sssnisreccmaneceoms 
IPhenacosaurus 2. eee 
Phyllobates............... 151; 1,.2) 4,592 
DOAETICIAS! -nccscsssedeneo iene 151, 5 
RIM OS DUN yde.terns surat 151, 7519 
LatIMASUS  Sasesecscosniceuni name 
tier 151, 2, 5, 6, 7, 9) 10 ye 
hb Fen oh nik: emeeneahen giane 151, 5, 9 
NUDICOLaA reece 161, 75 8;O7mu 


Occasional Papers of the Museum of Zoology 9 


sylvatica - tee Se ee 151, 7 

PELL AINANCAG! ccc. eee se 
151, 2, 4, 5, 6, 7, 9, 10, 11 
(ELA Fi eae ele act aT} Dy ae, 
Piyhodastylos 143, 3 
EIGN Fe. oka eo 143, 3 
Marin ee 143, 3 
puleher eee 143, 3 
iPhylomedusa. 2202 151, 2 
Physa spiculata ............ Eat LSD;: DS 
Pimephales notatus................ 138, 2, 3 
JO eS Se ener 136, 1 
BEG A cient rence 136, 1, 2 
[111 0 eA ee onee ee 136, 1, 2 
snethlegeze arated 136, 2 
Pisidium atlanticum .................. 135, 2 
1 Ea 152, 74 
dentiens cannarum.............. 135, 2 
obstructa 2 
orbicula 2 
ERIAMOP DIS’ otitis 1 
antrosus 2 
jordanensis E 
Dicarinatus ....... 1 
WMGHANENAIN) | Peso coco. 3 
circumlineatus......152, 71, 72, 74 
“Ed eta 1) (5 Ena ea a 135, 2 
TL EEDUTS LIE Sg Iie 135, 2 
Urls: 11 eee 152, 72, 74 
meridaensIB............00 152, 72, 74 

EQUI [IV | ee aes een oe 
oie 152, 71, 72, 73, 74, 111 
POTCATIMACUS niece cseessseensees 145, 1 
POTEAQENSIS 2... eecceeceeeneee 145, 1, 2 
SEU) ee 135, 2 
TOV ALONISIS 2 ection tensclcee 145, 1, 2 
ST] (EL (ge eg eh ee 145, 3 
U0) = ee 152, 72 
SUEIPCE CCE Ln eee eee 135, 2 
WETMIATIGT osecicossssssssessee 152, 72, 74 
VL IGLCNASY ren b sete sien 135, 2 
Pleurodema brachyop........... 152, 107 
Pineria bonairensig............... 152, 95 


Pleurodonte ineerta.................152, 107 


BApOTClay se ee 152, 108 
Lebire) | a Se eee lees, See ae 152, 107 

Poecilichthys coeruleua............. 138, 2 
WON eee ce et 138, 2 

Polylepiscus sp. 133, 2, 47 

Potamopyrgus 
parvulus.............. 

(PODS Meee eae eee 137, 28 
COMES Ae ee ee ee 137, 30 
corpulenta 137, 28 
dunkeri 137, 39, 40, 45 

CardOZI.........------ 137, 40, 43, 44 
Qumnkeri es raccans 137, 39, 40 
DOReZt en ccee 137, 39, 40 
POpaAyvanas ese 137, 45 
LY SOUR EPO coed a eS 137, 33 
gigantea... 137, 40, 42, 45 
fischeri............. 137, 40, 45 
STAUCOSEOMNA ns cececsscersnesoen 137, 36 
AViatic 137, 36, 37, 38 
glaucostoma ce 137, 38 
granadensis rugata........137, 36 
TCA eens 137, 40, 41, 46 
j Aeterna 137, 39 
popayana nn... 137, 45, 46 
CU RG Oa eed aes 137, 40 
POPAYV AMA... eeeeeennnn 137, 40, 46 
QUILENSIS See 137, 40, 45 
AMET qe ee ete 137, 35 
straminea.....137, 33, 35, 36, 38 
translucida........... 137, 31, 35, 41 
bejamensis |... 
ees 137, 31, 33, 34, 35 
major... 137, 32, 33, 35 
SANTAQUITENSIS neem 
pce tates a 137, 32, 33, 35 
translucida.........137, 31, 34, 35 
EY INILGUAIN eiceec 137, 31 

Proserpina swifti ...............137, 23 

Prostemmiulus heterops...133, 3, 15 

Pupa: tabeer 222 152, 75 
longvorigs. sae 152, 74, 116 
D1 fo isn he St 152, 98 


10 University of Michigan 


Pupisoma dioscoricola insigne...... 


BA Fra ee Pa Ee eee 135, 3 

PUPOIdes. careers: 152, 74, 116 
marginatus nitidulug............... 

Aero Us 137, 5; 152, 117 
IMOGUCTINY roe eats eee 137, 5 
simoni.......137, 5; 152, 75, 117 
SDl ee es 152 fo, lily 
Pycnotropis colombiensis-............... 
beet Pete 133, 2, 42, 44 
cylindroides ............ 133, 2, 43 
R 

Radelitiella tence ncncteees: 144, 1, 2 

dE efeitae |, coeeae repent a orate rete etree 151, 11 
WOU EICS oes iecerttstepconssete es 129, 10 

Rhacoma crossopetalum............ 152, 10 

Rhinocricus amblus........... ale eet!) 
ATDONCUS ne ae, 133, 3, 36 
BEEVIPES, 5.52252. 133, 2, 36 
hylophilus? 2s 133, 2, 36 
monilicornis ........... 133, 3, 36, 42 
DY. CNUS ies ere ec nec 133, 2, 38 

Toby PHODCSINTS - 2. wisconsin 133, 45 
amphelictus «0... 133, 3, 44 

Riccia wiOItAnS nese ene 138, 3 

Bomex brittamiea jesse. 138, 2 

S) 

Salasiella DrOWMi un... 135, 7 
guatemalensis .............. 135, 7, 8 
DINKIGV Ny aor nee eee 135, 7 
JOAQUINSS eee amare nual ane 135, 7 
MNAT GL ATIGACER, oaciectcesn en 135, 6 
Dulehella. eae ence 135, 7 

Salvelinus fontimalis................... 138, 3 

SED US Ltt ae eh 130, 4 

Scutellaria lateriflora................ 138, 2 

Semotilus atromaculatug........... 138, 2 

Siphonophora corynetes......133, 3, 8 
BPACINCEDB oy ecsimtnn: 133, 2, 5 
QTACIICOTNAS ........ccccceen So, Oy 
PLE 1 133, 3, 7, 9 
DEATSEL ce sitbiissscidices 150; 2), Os 7 
COMGtA: sii ennaaa 133, 3, 8 


Siphonotus parvus ©... 133, 3, 9 
PULPUTCUS nce nee 133, 9 
OLAS coe taseeanioe, cose eer 148, 1, 7 
pilineata: 2 ae 148, 7, 8 
DIMMNEAGIS | a racnseev eee 148, 8 
NOVINAC | pcttoeiek anor eee 148, 8 
WATICSALA leenansemeeee 148, 8 
Nomatochlora eee 130, 2 
Sparcaninm: (228 eee 138, 2 
Sphaerodactylus cinereus... 
Sah eee 182, 2, 30a 
ClESANS eee teces 132, 2, 3, 8, 9 
IMtermedius .......-..... 132, 2, 3, 9 
MOtAtUS | oie acon 132, 9 
Sputator 222i 132, 2 
Spiraxia. irrigwa; cee 135, 5 
shuttleworthi tc... 135, 5 
Spirostreptus atOPOrus cece 
sc erie on otter 133, 2, 21, 28 
Stemmatoiulus majov.............. 133, 10 
Stemmiulus craurus .......... 133, 2, 10 
drymophilus ...........-. 133, 3, 13 
labbanus ee 133, 3, 14 
TNA OL crete cree 133, 2, 10 
TUTHVENL 2.onsanccske 133; 2, a8 
BY) sf caciccsana aceon torminee 133, 2, 15 
Stenogyra octonoides.......... 152, 108 
SHEEKIA + AKOL be reagn eee 135, 3 
Stoastoma............-. 152, 34, 35, 111 
GOMINGENSIS) ssc 137, 22 
StOAStOMOPS sec iecccednc eee 
152, 35, 109, 111, 112, 114 
WOUKGTL Gonscansictnscteeee 152, 33 
Streptostyla cingulata.............. 135, 5 
Cylindraced si cchsnncees 135, 6 
CU WATOSIAN) ceeren ee 135, 5 
ATTIC UG) jot ote 135, 5, 6 
CQ WATASTATIA -essssersssserren 135, 5 
AUT OWA a ee 135, 5, 6 
shuttleworthi 0.0.0... 135, 5 
BUTTS Perneserares 1355-4), 0; .6s00 
VONUGIICOSA) Gites aurea 135, 5 
ERG VIB sce ccccerssecccie eee 7 
lattrei 6 
quirozi 6 


Occasional Papers of the Museum of Zoology ib 


Salle sacngsS.casesmes een 135, 5 
shuttleworthi .................. 135, 5, 6 
VGNLEICOSA aac iee 135, 5 
SUNT GS hea catinanenete nares 135, 5 
StreptOsty la rascces-cececcae: 135, 6 
Slcgined, bTeviswessccin arenes 135, 4 
PUACCMALCTSIS, ccrrrecsssneces 135, 3 
gyrata.........152, 18,°75, 110, 115 
MUUCTOS UNA recenrsrsnscesrineceee 135, 4 
DUC DICTISIS ie rrtesccnsncsiesatnnves 135, 4 
SUT LEN Alpe acces cerevecsaseccsesscatacsstec 135, 3, 4 
HAUT CT OSU Abe easerectacorttnnsosoqre 135, 4 
MOUS DLO SIS isc seccecssseecersseesce 135, 4 
T 

Theodoxus meleagris .............. 152, 32 
Thecadactylus rapicaudus .....143, 3 
Thysanophora ............... dogs 1 dts 
Camas) et creqamacticec 152, 79 
CONSPUTCACEL A eececeeeenesene 152, 79 
(Oh ialaaii 2 Ys Reaareercie pe eter 152, 109, 111 

hiahel| oye ale tay eee Oe en 
nae 152, 77, 78, 79, 106, 114 
CLIC eeeeseenenereenee 152, 78, 79 
TETUNS(CTE I aeetearetee eee reece 152, 79 
oO) SPAN eect i se Sa 135, 1 
Sth) 135, 1 
PROPRUNIN Eo ecstestiseeccetslteeerace 152, 79 
‘Sf oF UU ey, reece ere 152, 80 
Nyeatlcltiticldee ese one 152, 79, 114 
Tillandsia utriculata ................ 152, 21 


Trachelodesmus ..133, 53, 56, 59, 60 
ancylophor ..133, 2, 49, 59, 60 


AN GULACUS @ecesecceoeee ee 133, 2, 47 
constrictus ............ 133, 49, 50, 51 
LU DSTTE EY, ES eae eee ae eae 130, 2 
Tretioscincus bifasciatus ....143, 9 
UG Ge): a 152, 78 
VENEZUCIONSIS eeceeeeeseeeesee 152, 78 
Trichodiscina crinita ............. 152, 78 
Trichomorpha angulella ............... 
i Set A ses iy RE 133, 2, 66 
CUSeImMae ee 133, 2, 65 


NWI A at cette 133, 2, 64 
paurothrix —.._...._.. 133, 2, 66 
rugosella ............... 133, 2, 61, 63 
SCLOSION) asco case. 133, 2, 63 
tuberculosa ........ 133, 2, 60, 63 
rion 22.3.0 eet 133, 40, 42 
lumbricinus ............ 133, 3, 36, 42 
Propidodsetyiis 22 sla, al PAS) if 
Tropidonotus obliquus .............. 140, 1 
sipedon fasciatus .............. 140, 2 
iroschelivantd exqeesese cena 152, 111 
Truncatella bilabiata. .......... 152, 106 
Pudoracs ce 152, 16, 35, 41, 42, 
109, 113, 115, 116 
ATIPCNSIS 3e ee 137, 27 
UG ay ee 152, 49, 50, 51, 
56, 63, 111, 114 
aurantia. ......... 152, 45, 47, 48 
WasSSallenSiS: cee 
sha bea eet et 152, 47, 48, 61 
costata ..137, 3; 152, 45, 51, 53 
fossor .....152, 69, 110, 114, 115 
arubanay co eee 
eae 152, 67, 68, 70, 115 
Canashittensis) ..-----—-- 
ae 152, 26, 36, 67, 68, 69 
Gyerimensistess = 
. 152, 18, 54, 60, 65, 67, 68 
fOSSORE= 152, 36, 63, 65, 
66, 67, 68, 69, 70 
WEStPUNTENSIS eccrine 


152, 20, 66, 67, 68, 69, 70 
maculata, ...152, 36, 39, 42, 116 
megacheilos ............ 137, 2, 3, 27; 

152, 52, 57, 60, 62, 63, 64, 

65, 114, 115 


Kabrietensis: =... 
rts? 152, 14, 58, 59, 61 
TCO ACHOM OS sete tet tsa: 
oe 152, 55, 56, 58, 59, 61 
rondeklipensis ecco 
carne 152, 58, 59, 60, 61 
SPOR GS) NES OLY Lol Ses ao ieee 


12 University of Michigan 


MOBO ee 152, 45, 46, 50, 
53, 54, 111, 114, 115 
bullenensis ..152, 18, 47, 54 
grandiensis 
152, 47, 51, 52, 53, 54, 55 
MUSKUSI =e 152, 47, 51 
Pulsbryigence 152, 59, 62, 64, 
65, 113, 114, 115 

plicatala 4.22). 137, 23, 24, 
25, 26, 27; 152, 45, 46, 111 
Tupis ...152, 46, 52, 53, 111, 115 


MEWDPOItENSIS =... 
pe Ae eR 152, 47, 50, 113 
rupis.........152, 47, 49, 50, 113 
BUTEUILSS eee ee eco Nhe 152, 55 
WEPACGOT oe 152, 45 

williamsoni: —._--_.. 
Levee 137, 24, 26, 27; 152, 42 
Secand 137, 25, 27 
Turbo aurantius................... 152, 45, 46 
lab a) Se. eee 152, 42 
{U0 (2 eee reee eer ee eee 152, 98 


Typhlomanoleme ~ccrcccccccne-e 133, 19 
adapts <-e 133, 3, 19 
U 
Umbra lm 138, 2, 3, 4 
Uncancylus) sp. ie 135, 3 
Unio plicatulus.......__ 135, 1 
V 
Vaginula moreletie...... cece 135, 16 
x 
XONEn GUM ee tae 148, 1, 2 
calienten = ae 148, 1, 3, 4 
KANSCONG. a eee 148, 2 
CL PHOCSECUS) ec 120 7 
Z 
Zigwadesmus brunneus......133, 73, 74 
guiananusg............... 133, 3, 73, 74 
TNOGESTtUS ----svseseeccvseeeeeeeonees 133, 3, 74 
Zoogoneticus robustus......... 148, 4 


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